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Cell division [HTML]
Cell division
...ology View/Add Comments Table of Contents 1. The C. elegans embryo as a system to study cell division 2. A timeline of events ...

... events during the first mitotic division of the C. elegans embryo 3. Nuclear envelope structure and dynamics 4. Pronuclear mi...

...11. Acknowledgements 12. References Abstract The C. elegans embryo is a powerful model system for studying the mechanics of me...

...ristics that contribute to the usefulness of the C. elegans embryo for cell division studies. We provide a timeline for the fi...

Cell division : 1. The C. elegans embryo as a system to study cell division
...10.1895/wormbook.1.72.1 1. The C. elegans embryo as a system to study cell division The C. elegans embryo is a powerful model system for studying the mechanics of me...

...oocytes ( Figure 1 ). Since depletion relies on the rate of embryo production instead of protein half-life, the kinetics tend ...

... a protein required for cortical contractility in the early embryo ( Maddox et al., 2005 ). Serial dilutions of identically pr...

...l advantages contribute to the usefulness of the C. elegans embryo as a model system. Of particular importance is their rapid ...

Cell division : 2. A timeline of events during the first mitotic division of the C. elegans embryo
..., the nuclear-centrosome complex moves to the center of the embryo and rotates to align with the long axis of the embryo ( Albertson, 1984 ; Hyman and White, 1987 ). The miotitc sp...

...etry see Asymmetric cell division and axis formation in the embryo ). Figure 2. Nuclear envelope dynamics in the C. elegans embryo.  (Left column) Schematics illustrate the major featur...

... events during the first mitotic division of the C. elegans embryo Due to its accessibility to RNAi-based molecular perturbati...

...ed by a small acentriolar meiotic spindle that forms in the embryo anterior. During anaphase of meiosis I and again in meiosis...

Cell division : 3. Nuclear envelope structure and dynamics
...ssembly during the first mitotic division of the C. elegans embryo are illustrated in Figure 3 . LMN-1 leaves the nuclear enve...

...ion). Figure 3. Nuclear envelope dynamics in the C. elegans embryo.  Figure courtesy of Vincent Galy. (A) Schematics illu...

Cell division : 4. Pronuclear migration
...sp;Pronuclear migration The site of sperm entry defines the embryo posterior ( Goldstein and Hird, 1996 ). As the pronuclei be...

... rounds of meiotic chromosome segregation, typically in the embryo anterior. The two pronuclei migrate towards each other coin...

...nt of the sperm pronucleus away from the cortex towards the embryo center ( Albertson, 1984 ; O'Connell et al., 2000 ). Initia...

....5 μm/min till it meets the oocyte pronucleus near the embryo center (~7 μm). Figure 4. Kinetics of pronuclear migra...

Cell division : 5. Centrosome assembly and duplication
...es, one inherited from the sperm and one that formed in the embryo cytoplasm. As the embryo enters mitosis, the amount of PCM around the centrioles and...

...r, 2003 ). Figure 5. Mitotic kinetochores in the C. elegans embryo.  (A) Schematic of the first cycle of centrosome dupli...

...ntrosomes recruit additional pericentriolar material as the embryo enters mitosis in a process called centrosome maturation, i...

...pronuclear migration fails and a short spindle forms in the embryo posterior. Matthews, 1998 ; Le Bot, 2003 ; Srayko, 2003 ; B...

Cell division : 7. Kinetochore assembly
...., 2004 ). Figure 6. Mitotic kinetochores in the C. elegans embryo.  (A) Images comparing the localized kinetochores in a...

...ric chromosomes to the diffuse kinetochores in a C. elegans embryo with holocentric chromosomes. (B) Schematic illustrating th...

Cell division : 8. Assembly of the mitotic spindle
...e mechanisms that regulate spindle length in the C. elegans embryo are largely independent of bipolarity. Experiments in which...

...hat normally asymmetrically position the spindle within the embryo ( Oegema et al., 2001 ). The abrupt separation of the spind...

Cell division : 10. Cytokinesis
...at are known to contribute to cytokinesis in the C. elegans embryo can be partitioned into three classes ( Table 5 ). The firs...

... the only C. elegans profilin that is required in the early embryo; depleted embryos exhibit defects in furrow ingression and ...

...that appears to have a role in cortical events in the early embryo; localizes to the contractile ring, as well as to other cor...

...t are able to form cortical ruffles that concentrate in the embryo anterior during polarity establishment and a pseudocleavage...


Asymmetric cell division and axis formation in the embryo [HTML]
Asymmetric cell division and axis formation in the embryo
...y an important role in generating diversity during metazoan development. In the early C. elegans embryo, a series of asymmetric divisions are crucial for establish...

...k.1.30.1 Asymmetric cell division and axis formation in the embryo Download PDF version Asymmetric cell division and axis form...

... version Asymmetric cell division and axis formation in the embryo * Pierre Gönczy § Swiss Institute for Experimenta...

...nts Table of Contents 1. Introduction 2. The one-cell stage embryo: establishment of the anterior-posterior axis 2.1. Breaking...

Asymmetric cell division and axis formation in the embryo: 1. Introduction
...etric divisions are crucial for generating diversity during development because they yield daughter cells that differ in fate. In C...

... four-cell stage, with EMS defining the ventral side of the embryo. Third, the left-right (LR) axis is detectable at the trans...

...hters of AB ( ABal and ABpl ) defining the left side of the embryo. Because the position of the mitotic spindle in animal cell...

...re detailed coverage of the underlying data. Movie 1.  Embryo carrying a GFP-histone2B fusion protein (Strome et al., 200...

Asymmetric cell division and axis formation in the embryo: 2. The one-cell stage embryo: establishment of the anterior-posterior axis
...10.1895/wormbook.1.30.1 2. The one-cell stage embryo: establishment of the anterior-posterior axis 2.1. Bre...

...g the sequence of events that result in polarization of the embryo along the AP axis. Consistent with this view, proper AP pol...

...twork ( Munro et al., 2004 ). Movie 3.  One-cell stage embryo carrying a GFP-NMY-2 fusion protein (Munro et al., 2004) im...

...ovie is displayed ~150 times faster than actual events. The embryo is approximately 50 μ m long and anterior is to the le...

Asymmetric cell division and axis formation in the embryo: 3. The one-cell stage embryo: spindle positioning
...10.1895/wormbook.1.30.1 3. The one-cell stage embryo: spindle positioning 3.1. Cortical forces position cen...

...s the male pronucleus and its associated centrosomes at the embryo's posterior, the centrosomes and pronuclei move as a unit t...

...r, the centrosomes and pronuclei move as a unit towards the embryo's center during prophase (centration). The unit also rotate...

... polarity mediators and cell fate determinants in the early embryo.  P granules: black discs; cytoplasmic POS-1, MEX-1, a...

Asymmetric cell division and axis formation in the embryo: 4. The two-cell stage embryo: formation of the dorsal/ventral axis
...10.1895/wormbook.1.30.1 4. The two-cell stage embryo: formation of the dorsal/ventral axis The daughters of the ...

...1 . GPR-1/2 and LET-99 are not shown: in the early two-cell embryo, they are initially enriched in patterns similar to that se...

...P 2 cell contact. After asymmetric division of the two-cell embryo, localized cell fate determinants lead to unique fates for ...

... P 2 and EMS , with EMS defining the ventral portion of the embryo ( Figure 4 C). Further, the second division spindle positio...

Asymmetric cell division and axis formation in the embryo: 5. The four-cell stage embryo: formation of the left/right axis
...10.1895/wormbook.1.30.1 5. The four-cell stage embryo: formation of the left/right axis At third cleavage, the le...

...ay is responding to an asymmetry established earlier during development. After the ABa and ABp cells divide, EMS divides asymmetric...

...ocalized to specific cells or subsets of cells in the early embryo ( Figure 4 ). These determinants then interact to instruct ...

...al program which will ultimately result in the making of an embryo ready to hatch. ...

Asymmetric cell division and axis formation in the embryo: 6. Evolutionarily conserved themes
...ed themes Many of the findings made in the early C. elegans embryo and that were briefly reviewed above have had important bea...

...wise, the specification of the other axes of the C. elegans embryo, as well as that of particular cell fates, utilize highly c...

...ways (see Wnt signaling , Notch signaling in the C. elegans embryo and LIN-12/Notch signaling in C. elegans ). Thus, just as t...

... elegans ). Thus, just as the study of the early C. elegans embryo provided key insight into developmental processes, studies ...


Notch signaling in the C. elegans embryo [HTML]
Notch signaling in the C. elegans embryo
...ml 10.1895/wormbook.1.4.1 Notch signaling in the C. elegans embryo Download PDF version Notch signaling in the C. elegans embryo * James R. Priess § Fred Hutchinson Cancer Research Ce...

...tors are major targets of Notch signaling in the C. elegans embryo, and are distantly related to HES proteins that are targets...

Notch signaling in the C. elegans embryo : 1. Introduction
...otch signaling controls mesoderm induction during embryonic development, and controls germ cell mitosis during postembryonic development (see below and Austin and Kimble, 1987 ). The general mecha...

...gnaling pathway plays a central role in patterning metazoan development, and is used in remarkably diverse cell fate decisions (for...

...have been documented throughout embryonic and postembryonic development of C. elegans . For example, Notch signaling controls mesod...

... LAG-1 /Su(H). A principal target of Notch signaling in the embryo is the ref-1 gene family, consisting of ref-1 , hlh-25 , hl...

Notch signaling in the C. elegans embryo : 2. The first and second interactions in the AB lineage
...ntact with the ligand-expressing P 2 cell; thus in a 4-cell embryo GLP-1 /Notch is activated in ABp , but not in ABa ( Figure ...

...eres at the 2-cell, 4-cell, and 12-cell stages; the 12-cell embryo is a ventral view with the AB descendants splayed to demons...

... to be refractive to the second Notch interaction in normal development, and can respond only if the first Notch interaction is blo...

...on of a cell is considered here as its 'fate'. In wild-type development, the eight AB descendants have unique fates that can be rep...

Notch signaling in the C. elegans embryo : 3. Notch-dependent/Notch-independent gene regulation
...to induce expression of PHA-4 , and thus promote mesodermal development, has not yet been identified. ...

Notch signaling in the C. elegans embryo : 4. The third AB Notch interaction: Formation of the bilaterally symmetrical head
...on: Formation of the bilaterally symmetrical head In normal development, an ABp descendant called ABplaaa and an ABa descendant cal...

...first evidence that cell interactions were involved in head development came from the finding that lin-12 glp-1 double mutants had ...

...hat it is not influenced by cell interactions . However the development of the left head precursor is altered markedly when precurs...

...together with GLP-1 /Notch cause similar defects in ABplaaa development, indicating that the fate of the left precursor is specifie...

Notch signaling in the C. elegans embryo : 5. The fourth AB Notch interaction: specification of the excretory cell
... Schnabel, 1995a ; Moskowitz and Rothman, 1996 ). In normal development one of the ABplpapp descendants produces the excretory cell...

Notch signaling in the C. elegans embryo : 6. A combinatorial Notch code for AB specification
...). TBX-37 , -38 contribute to these primary fates in normal development, and a failure in the first interaction causes misexpressio...

...h an unknown target of Notch signaling to induce pharyngeal development ( Good et al., 2004 ). The third and fourth interactions co...

Notch signaling in the C. elegans embryo : 7. Intersection of the Notch pathway and the POP-1 polarity pathway
...lls in C. elegans , and suggested that cells throughout the embryo somehow recognized a common anterior/posterior axis of pola...

...he MS blastomere is positioned in the middle of a wild-type embryo, and the MS descendant born from the cleavage pattern poste...

... ectopic-MS-like cells in the anterior and posterior of the embryo, respectively ( Mello et al., 1992 ). Remarkably, the anter...

...e posterior fate. POP-1 asymmetry is generated in the early embryo by a non-canonical Wnt pathway involving the ligand MOM-2 /...

Notch signaling in the C. elegans embryo : 8. Notch interactions in later AB descendants
... inductive focus of Notch signaling on the left side of the embryo. Schematic diagram of a cross section through an embryo at successive time points. MS descendants are indicated wit...

...n-12 glp-1 double mutant embryos have several defects in AB development beyond those described above, indicating that there are mul...

...ranscription factor PAL-1 appears to be required for rectal development ( Edgar et al., 2001 ) and appears to be a direct target of...

...dants appear to be one of the main signaling centers in the embryo. Figure 7.  MSap descendants provide an inductive focu...

Notch signaling in the C. elegans embryo : 9. Notch interactions in P1 descendants
...cendants express GLP-1 /Notch or LIN-12 /Notch in wild-type development (J. Priess, unpublished). However, Notch interactions thus ...

...re intestine ( Hermann et al., 2000 ). Stages of intestinal development are designated as E2, E4, etc. to indicate the number of E ...

Notch signaling in the C. elegans embryo : 11. Perspectives
...oan signaling pathways, the Notch pathway, functions during development. Our current view of the Notch network in the embryo will undoubtedly become more elaborate as additional intera...

...ectively serves to replicate a Notch-independent pattern of development in additional parts of the embryo. In future studies, it will be interesting in to see whethe...

...o be the major effector of Notch-mediated repression in the embryo, and analysis of this gene family should provide insight in...


Translational control of maternal RNAs [HTML]
Translational control of maternal RNAs
...ational control of the Notch signaling pathway in the early embryo 3.2. Control of pal-1 in the embryo 3.3. Regulation of maternal mRNAs and germ cell specificati...

...precise temporal and spatial patterns of translation in the embryo. In the embryo, cell polarity factors control the localization of translat...

... mRNAs in the germ line 3. Control of maternal mRNAs in the embryo 3.1. Translational control of the Notch signaling pathway i...

...lation of maternal mRNAs and germ cell specification in the embryo 4. Conclusions and perspectives 5. Acknowledgements 6. Refe...

Translational control of maternal RNAs: 1. Introduction
...10.1895/wormbook.1.34.1 1. Introduction Early development of C. elegans embryos requires the localization of maternal...

...ve been described for localizing maternal regulators in the embryo; directed movement or local trapping of maternal proteins, ...

...NAs (see Asymmetric cell division and axis formation in the embryo ). This chapter reviews the translational regulation of mat...

...controlled, and how their control is connected to germ cell development and early embryonic polarity. We define "translational cont...

Translational control of maternal RNAs: 2. Control of maternal mRNA translation begins in the hermaphrodite germ line
...w specific patterns of protein accumulation to arise in the embryo. We limit our discussion to mRNAs that function in the embryo. For discussion of the regulation of other classes of germl...

...ermaphrodite germ line Most maternal mRNAs destined for the embryo are transcribed by germ cell nuclei in mitosis or early sta...

...on of many maternal mRNAs is repressed during female gamete development ( Table 1 ). This tight regulation is likely necessary to p...

...ion is likely necessary to prevent interference with oocyte development and is essential to allow specific patterns of protein accu...

Translational control of maternal RNAs: 3. Control of maternal mRNAs in the embryo
...ational control of the Notch signaling pathway in the early embryo In the early C. elegans embryo, Notch signaling between posterior and anterior blastomeres...

...95/wormbook.1.34.1 3. Control of maternal mRNAs in the embryo Maternal RNAs that have been silenced in the germ line are ...

...ated in specific spatial and temporal patterns in the early embryo ( Figure 1 ; Table 1 ). Because early cell cycles are rapid...

...ulators are localized to specific blastomeres of the 4-cell embryo.  APX-1/Delta is localized to the posterior blastomere...

Translational control of maternal RNAs: 4. Conclusions and perspectives
...ently to manage message specific expression patterns in the embryo. Second, regulated maternal mRNAs generally contain multipl...

...hanism, which might be poised for rapid derepression in the embryo ( Mootz et al., 2004 ). This mechanism may be similar to mi...

...in the germ line but translated in opposing patterns in the embryo? What is the composition of translational control complexes...

...ontrol activities spatially and temporally regulated in the embryo? These questions will provide the fuel for exciting researc...


Epidermal morphogenesis [HTML]
Epidermal morphogenesis
... 8. References Abstract The morphogenesis of the C. elegans embryo is largely controlled by the development of the epidermis, also known as the hypodermis, a single ep...

...pe, epidermal cells are generated on the dorsal side of the embryo among the progeny of four early embryonic blastomeres. Spec...

...pidermal sheet undergoes epiboly to enclose the rest of the embryo in a process known as ventral enclosure; this movement requ...

Epidermal morphogenesis : 1. Introduction
...5/wormbook.1.35.1 1. Introduction Morphogenesis is the development of form, of tissues, of organs, and of organisms. This chap...

...major morphogenetic processes of the mid-to-late C. elegans embryo, focusing on the epidermis and its interactions with underl...

...has not been fully elucidated. In this review we follow the development of the epidermis during embryogenesis, focusing on processe...

...lular matrix, including the cuticle, in embryonic epidermal development. ...

Epidermal morphogenesis : 2. Formation and specification of epidermal cells
...ns, 220-240 minutes after first cleavage; at this stage the embryo is composed of ~365 cells. The major epidermal cells are de...

...l precursors are located on the dorsal surface of the early embryo, and thus the major epidermal cells arise as a group on thi...

Epidermal morphogenesis : 3. Movements of ventral neuroblasts
...or later enclosure movements of the epidermis, and so their development is considered here. Ventral neuroblasts themselves undergo ...

...dy muscles ingress on the ventral surface of the developing embryo (see Gastrulation in C. elegans ). These ingression movemen...

... period of epidermal enclosure when the ventral side of the embryo first becomes visibly concave. Note that unequal growth alo...

...at unequal growth along the anteroposterior axis forces the embryo to turn within the eggshell between 350 and 400 minutes so ...

Epidermal morphogenesis : 4. Intercalation of dorsal epidermal cells
...s make contacts with seam cells on the opposite side of the embryo ( Sulston et al., 1983 ; Williams-Masson et al., 1998 ; Fig...

....   Movie 2.  Dorsal intercalation in a wild-type embryo visualized using Nomarski microscopy ( Williams-Masson et a...

....   Movie 3.  Dorsal intercalation in a wild-type embryo visualized using a dlg-1 ::gfp translational fusion ( Koppe...

... shown in white. Dorsal cells born on the right side of the embryo are depicted in a lighter color than those born on the left...

Epidermal morphogenesis : 5. Ventral enclosure
...reme anterior epidermis then completes the enclosure of the embryo by a sheet of epidermis ( Figure 4 ).   Figure 4. Epid...

...(D).   Movie 7.  Ventral enclosure in a wild-type embryo visualized using a dlg-1 p::gfp transcriptional reporter ( ...

...g dorsal intercalation, corresponding to Figure 3B . (B) An embryo expressing an lbp-1 ::gfp transcription reporter to mark ep...

...ring ventral enclosure, corresponding to Figure 4B . (E) An embryo expressing a dlg-1 ::gfp transcriptional reporter ( Fireste...

Epidermal morphogenesis : 6. Embryonic elongation
... epidermis is enclosed, elongation converts the bean-shaped embryo into the elongated shape of the worm ( Movie 9 ). In our di...

...92 ), the pharynx and intestine may not be required for the embryo to elongate. However, at least one internal tissue (body mu...

... epidermis.   Movie 9.  Elongation in a wild-type embryo imaged using Nomarski microscopy. Frames were collected at ...

...omplete by 600 minutes ( Figure 1 ). During elongation, the embryo reduces its circumference by a factor of three and increase...


Gastrulation in C. elegans [HTML]
Gastrulation in C. elegans
...he process by which the germ layers become positioned in an embryo. C. elegans gastrulation serves as a model for studying the...

Gastrulation in C. elegans: 1. Summary of gastrulation events
...mall. Despite this, gastrulation plays an essential role in development, internalizing endodermal, mesodermal, and germ-line precur...

...s of cells separate from one another in the interior of the embryo. Cells acquire an apical-basal polarity that is important f...

...rds each other, displacing Ea and Ep into the center of the embryo. Myosin accumulates at the apical surfaces of cells as they...

...Embryological variation during nematode development . ...

Gastrulation in C. elegans: 2. Polarization of gastrulating cells
...n par(ZF1) embryos.  (A,C) Surface view of a wild-type embryo (A) or an embryo where PAR-6 has been removed by fusion to the ZF1 domain (C...

...al view of the interior region of a 26-cell stage wild-type embryo (B) or par-6(ZF1) embryo (D). A space (arrow) has formed between lateral cell surfac...

...dermal precursors, Ea and Ep , move from the surface of the embryo into a small interior cavity called the blastocoel ( Sulsto...

...l ( Sulston et al., 1983 ). Prior to the 26-cell stage, the embryo is organized as a hull of cells one cell in radius that is ...

Gastrulation in C. elegans: 3. Mechanisms of cell ingression
...ments. (A-D) Time-lapse images of gastrulation in an intact embryo (same embryo as Figure 1 C), (E-H) in a devitellinized embryo, and (I-P) in P 1 isolates. Variation of starting orientati...

...traints or a necessary microenvironment to the gastrulating embryo, the eggshell and vitelline envelope were removed ( Lee and...

...panel shows a ventral view, with Ea and Ep sinking into the embryo. Embryos are oriented anterior to the left. Figure 5. Gastr...

...bryos.  (A, a-c) Ea/p (asterisks) position in the same embryo filmed at the 24-cell stage (a), 28-cell stage (b) and 46-c...

Gastrulation in C. elegans: 4. Patterning cell ingressions
...s ). Hypodermal cells are born on the dorsal surface of the embryo and migrate ventrally to encase the embryo in skin. Figure 9. Cell ingressions during gastrulation.&nb...

...proximately 90 minutes after the first cell division of the embryo. After the endodermal cells have completed ingression, meso...

...astrulation. (B) Position of cell nuclei in a 26-cell stage embryo, lateral perspective, at the onset of gastrulation. Nuclei ...


RNA-binding proteins [HTML]
RNA-binding proteins
...ents 1. Overview 2. RBPs function during germline and early embryo development 3. RBPs function in somatic development 4. RNA targets of RBPs 5. RNA binding specificity of RBPs 6...

... many RNA-binding proteins (RBPs) with diverse functions in development, indicative of extensive layers of post-transcriptional con...

...characterized mutant phenotypes in the germ line, the early embryo, or in somatic tissues. We also discuss the identification ...

...ant first step to understand how an RBP controls C. elegans development. It is likely that most RBPs regulate multiple RNA targets....

RNA-binding proteins : 1. Overview
...n identified as essential factors during germline and early embryo development and RBPs with essential functions in the development of somatic tissues, including neuron, muscle, hypodermis, a...

...argets 1 References RBPs function during germline and early embryo development gld-1 Maxi-KH Female meiotic prophase progression, germline...

...gld-3 KH-like Gemline sex determination, germline survival, embryo development Translational activator? Unknown Eckmann et al., 2002 ; Wan...

...2000 mog-1 /-4/-5 DEAD/ DEAH box Gemline sex determination, embryo development RNA helicase Unknown Puoti and Kimble, 1999 ; Puoti and Kim...

RNA-binding proteins : 2. RBPs function during germline and early embryo development
...book.1.79.1 2. RBPs function during germline and early embryo development Many RBPs have essential functions during late germline and...

...BPs have essential functions during late germline and early embryo development ( Table 1 ) because post-transcriptional control of materna...

... ) have also been implicated in regulating aspects of early embryo development. These RBPs likely regulate the temporal and spatial patter...

...ermline sex determination and the proliferation vs. meiotic development decision (stem cell maintenance) rely heavily on post-trans...

RNA-binding proteins : 3. RBPs function in somatic development
... Table 1 ). Unlike RBPs that function in germline and early embryo development, several RBPs essential for somatic development are proposed to act as splicing factors, presumably regulat...

...10.1895/wormbook.1.79.1 3. RBPs function in somatic development Post-transcriptional control is also important in somatic development as a number of RBPs that have somatic tissue-specific mutan...

...dependent regulation occur during both germline and somatic development and recent discoveries of RBPs functioning in RNAi and miRN...

...this new mode of post-transcriptional control in C. elegans development. ...

RNA-binding proteins : 4. RNA targets of RBPs
...that a comprehensive understanding of how such RBPs control development requires the identification of many or all of their mRNA ta...

RNA-binding proteins : 5. RNA binding specificity of RBPs
... questions to understand the function of RBPs in C. elegans development is how RBPs distinguish their targets from non-targets in v...

RNA-binding proteins : 6. Closing remarks
... elegans genome encodes many RBPs with diverse functions in development, revealing a large layer of post-transcriptional control of...

... comprehensive understanding of how RBPs control C. elegans development. With multiple RNA targets, specific features that distingu...

... coordinate control of their RNA targets, thereby affecting development in a concerted fashion. ...


Methods in cell biology [HTML]
Methods in cell biology : 1. Introduction
...genetics ( Brenner, 1974 ; Jorgensen and Mango, 2002 ), the development of fluorescent protein tags ( Chalfie et al., 1994 ; Yang e...

Methods in cell biology : 2. Visualizing cells and their components
.... Transfer blocks to whatever mold you will use to hold the embryo and try your best to orient embryo for easiest trimming and cutting. 3 days Araldite @60°C...

...larvae seem to fix better than older stages. Interestingly, embryo fixation using standard methods is fairly difficult, thus, ...

...g and Bob Horvitz). Figure 10.  TEM section through an embryo fixed using HPF (Rick Fetter and Cori Bargmann). Because fi...

...ould be performed with a tissue rotator. In 100% resin, the embryo/worm plus E.coli /yeast pancake usually comes out of the ho...

Methods in cell biology : 3. Protein-protein and protein-DNA interactions
...fer, resuspend in 1 mL of homogenization buffer per gram of embryo and store in − 80°C freezer. Embryo Lysates All steps are performed at 4°C. Thaw 10–1...

... shift assays (EMSA). Protocol 16: Immunoprecipitation from embryo lysates (Ray Chan and Barbara Meyer) Worm Liquid Culture: S...

...s. Immunoprecipitation All steps performed at 4°C. Thaw embryo lysates on ice. Incubate lysates with Protein A Sepharaose ...

... of affinity purified antibodies with 3 mg total protein of embryo lysates for 2 hrs. Bring the final volume up to 1.0 to 1.4 ...

Methods in cell biology : 4. Specific methods in C. elegans cell biology
...soma and germ line. Figure 13.  DIC image of wild-type embryo at comma stage of development. Arrows point to the round, button-like corpses (Julia Hatz...

...ing at germ cells, intestinal cells, and cells in the early embryo. A variety of protocols related to chromatin visualization ...

...t be better for embryos. This protocol was not designed for embryo staining, although embryos do sometimes stain with antibody...

...al volume of fixative (4% PFA, 1× sperm salts) to the embryo pellet. Pipette 10 μL of embryos to each slide and cov...

Methods in cell biology : 5. Embryonic cell culture
...o generate GFP-labeled cells that emerge later in embryonic development (e.g. unc-4 ::GFP in DA motor neurons) (brown arrows). 4. G...

...re-pellet to remove the sucrose. Preparation of dissociated embryo cells After egg isolation, all subsequent steps should be c...

...out in a laminar flow hood under sterile conditions. Remove embryo eggshells by incubating approximately 50–200 μl ...

...profiles of specific cell populations ( Fox et al., 2005 ). Embryo isolation Synchronize worms by allowing them to exhaust the...


Cell fusion [HTML]
Cell fusion
...erences Abstract Selective cell fusion is a natural part of development. It is found in sexually reproducing organisms that require...

...nd stem cells. Cell fusion is particularly important in the development of C. elegans: in addition to 300 sperm and oocytes that fu...

...zation, 300 of the 1090 somatic cells born, fuse throughout development. Studies of cell fusion in C. elegans have shown that altho...

...he lateral epidermal cells that normally do not fuse in the embryo will fuse causing embryonic lethality. And if either the Ho...

Cell fusion: 1. Introduction
...; see Figure 1 ; Wormatlas , Epidermal morphogenesis , Male development and Vulval development ). These cells are formed during embryonic and larval development in a genetically programmed process. Syncytiogenesis culmin...

...ia are generated by the fusion of mononucleate cells during development. The largest syncytium is hyp7 which is generated in the embryo during morphogenesis ( Figure 1 ; Podbilewicz and White, 19...

... eye and stem cells, although most cells do not fuse during development. ( Baroux et al., 2004 ; Chen and Olson, 2004 ; Cross et al...

...ed for fusion (gray color and dashed lines) fuse throughout development forming different syncytia (colored). White marks unfused c...

Cell fusion: 2. Cell biology of plasma membrane fusion
...e in the understanding of developmental cell fusion was the development of a system to reproducibly observe ongoing cell fusions wi...

Cell fusion: 5. Regulation of cell fusion
...orphogenesis , Hermaphrodite cell-fate specification , Male development , and Vulval development ). 5.2.1. Hox genes protect specific cells from fusion...

...te in embryonic elongation but do not fuse in the wild-type embryo. However, during postembryonic development the ectoblastic seams have dynamic cell behaviors essential...

...f cells that fuse forming the hyp6 and hyp7 syncytia as the embryo elongates ( Podbilewicz and White, 1994 ; Priess and Hirsh,...

...ate neurons or maintain the seam cells during postembryonic development. 3) Two rows of ventral P blast cells that do not fuse in t...

Cell fusion: 6. Open questions
...appear to precisely regulate eff-1 transcription throughout development. It is expected that numerous posttranslational regulators ...

...g of cytoplasmic contents.  eff-1( hy21 ) ; hsp::EFF-1 embryo expressing ajm-1 ::gfp and eff-1p::gfp . Following heat sho...

...on? How are dorsal hypodermal cell fusions regulated in the embryo? ...

Cell fusion: 7. Implications of cell fusion research beyond worms
... become differentiated. Thus, defects in cell fusion during development may result in cancerous growth in mammals ( Duelli and Laze...


Vulval development [HTML]
Vulval development
...vulvaldev.html 10.1895/wormbook.1.6.1 Vulval development Download PDF version Vulval development * Paul W. Sternberg § Howard Hughes Medical Institute ...

...ents Table of Contents 1. Introduction 1.1. Steps in vulval development 1.2. Phenotypes of vulval development mutants 1.3. Note on phenotypes 2. Generation of vulval pre...

...tes 3.2. Commitment to fates 3.3. The cell cycle and vulval development 4. Induction of the vulva by the anchor cell 4.1. The ancho...

...T signaling and induction 5. Physiological inputs to vulval development 5.1. G protein regulation 5.2. Zinc regulation 6. Negative ...

Vulval development: 1. Introduction
...erves as a paradigm for organogenesis. In particular, vulva development represents a well-understood case in which invariant development arises from multiple cell-cell interactions. It is also a s...

... ) and Notch ( LIN-12 ) pathways. 1.1. Steps in vulval development Vulval development is conveniently thought of as occurring in five steps ( Fig...

...ite molts to adulthood. In addition, two aspects of uterine development are crucial to the development of the vulva. (A) Generation of the anchor cell. The anchor...

...nchor cell ultimately fuses with the utse. Figure 1. Vulval development is a multi-step process.  It involves the coordinated development of the uterine (green, top) and ventral epithelia (purple, ...

Vulval development: 2. Generation of vulval precursor cells
...e relatively subtle, but very intriguing in light of vulval development in other nematode species (see Evolution of development in nematodes closely related to C. elegans ). ...

...clei migrate into the ventral cord during the L1 stage. The development of the vulva thus depends upon the generation of the Pn.p c...

... redivivus ( Sternberg and Horvitz, 1982 ; see Evolution of development in nematodes closely related to C. elegans ), suggesting th...

...d UNC-62 are co-factors of LIN-39 and as such affect vulval development ( Yang et al., 2005 ). Two redundant GATA factors, encoded ...

Vulval development: 3. Overview of VPC 1°-2°-3° pattern formation
...e to respond to LIN-3 . 3.3. The cell cycle and vulval development Vulva development involves cell cycle in two distinct ways. Cell cycle regula...

... indicated. Wnt signaling plays several roles during vulval development, including specification of VPC competence and polarity of ...

....2. Commitment to fates An open question in C. elegans development concerns fate commitment. I use specification to mean a cel...

...easonable set of perturbations). The rapidity of C. elegans development, and the lack of long-term culture have precluded extensive...

Vulval development: 4. Induction of the vulva by the anchor cell
...ts ancestors prior to the L3 stage completely blocks vulval development, and the VPCs have 3 ° fates (3 ° -3 ° -3 °...

...ght be some that are physiologically relevant during vulval development. Mediator complex proteins, which link site-specific transc...

...on factors to RNA polymerase, also are necessary for vulval development. lin-25 and sur-2 ( mdt-23 ) are defective in vulval induct...

...ction and also have defects earlier and later during vulval development ( Ferguson et al., 1987 ; Tuck and Greenwald, 1995 ; Singh ...

Vulval development: 5. Physiological inputs to vulval development
....1895/wormbook.1.6.1 5. Physiological inputs to vulval development The invariance of vulval development is not only due to the growth of C. elegans under standardi...

...er standardized laboratory conditions; however, when vulval development is partly compromised, a number of physiological effects ar...

...at neuronal and muscle excitation can somehow affect vulval development. The positive signal from Gq acts in parallel to or downstr...

...rotein coupled receptor, SRA-13 , acts negatively on vulval development via the G protein GPA-5 ( Battu et al., 2003 ). Starvation ...

Vulval development: 6. Negative regulation of induction
...tants defective in only A, or in only B, have normal vulval development. Molecular cloning revealed that many Class B synMuv genes ...

Vulval development: 8. Coupling of LET-23 and LIN-12 signaling
... Berset et al., 2001 ; Yoo et al., 2004 ). C. elegans vulva development was the first case where the interplay of growth factor rec...

Vulval development: 10. The vulval-uterine connection
...11 and cog-2 , two transcription factors necessary for utse development, including fusion of the anchor cell with utse. Model adapt...

Vulval development: 11. Morphogenesis
... of neurons and muscles The vulval epithelium organizes the development of the muscles and neurons that mediate egg-laying ( Li and...

...ucts expressed in subsets of vulval cells direct details of development of individual neurons. bam-2 is necessary for axonal branch...

Vulval development: 12. Concluding remarks
...larly, our understanding of the control of timing of vulval development is also incomplete, and the relationship of the cell cycle ...

...e limitations, the extensive knowledge of C. elegans vulval development makes it an interesting case for computational modeling stu...

Vulval development: 13. Acknowledgments
...y colleagues over the past 25 years who have studied vulval development or at least been willing to discuss it, especially Bob Horv...

.... I am grateful to the following sources that funded vulval development research in my laboratory: Searle Foundation (1988-1991); N...


The C. elegans pharynx: a model for organogenesis [HTML]
The C. elegans pharynx: a model for organogenesis
...ments Table of Contents 1. The pharynx as a model for organ development 2. Anatomy of the pharynx 3. Early control of pharyngeal development-the ABa lineage 4. Early control of pharyngeal development-the EMS lineage 5. The transition from maternal to zygotic ...

...e specification and epithelial morphogenesis during pharynx development. Maternally-supplied gene products function prior to gastru...

The C. elegans pharynx: a model for organogenesis : 1. The pharynx as a model for organ development
...nt embryos at the level of individual cells. Third, pharynx development is robust. Embryos with abnormal development in other tissues can still produce a well-differentiated ph...

...5/wormbook.1.129.1 1. The pharynx as a model for organ development Four characteristics of the C. elegans pharynx make it a po...

...till produce a well-differentiated pharynx. For example, an embryo that cannot undergo normal morphogenesis arrests as a ball ...

...FoxA proteins, and, like pha-4 , FoxA2 is essential for gut development in all organisms studied to date ( Carlsson and Mahlapuu, 2...

The C. elegans pharynx: a model for organogenesis : 3. Early control of pharyngeal development-the ABa lineage
...iess, 2005 ). Table 1. Summary of genes involved in pharynx development. Genes implicated in pharynx development are listed from earliest stages of specification to later e...

...lar biology section of WormBook. Figure 4. Early pharyngeal development.  Features of pharyngeal development from the 4-cell stage to the 28-cell stage. This period is ...

...0.1895/wormbook.1.129.1 3. Early control of pharyngeal development-the ABa lineage The pharynx is generated polyclonally durin...

...7 ; Priess and Thomson, 1987 ). glp-1 RNA is donated to the embryo by the mother (i.e., maternally) and is selectively transla...

The C. elegans pharynx: a model for organogenesis : 4. Early control of pharyngeal development-the EMS lineage
...0.1895/wormbook.1.129.1 4. Early control of pharyngeal development-the EMS lineage The EMS pathway of pharyngeal development does not require glp-1 signals (see Figure 4 ; Good et al.,...

...requirement for Tbox factors for both ABa and MS pharyngeal development. A second target of SKN-1 within MS are ref-1 family member...

The C. elegans pharynx: a model for organogenesis : 5. The transition from maternal to zygotic control
...nd the MED factors initiate the zygotic phase of pharyngeal development. The phenotypes associated with the zygotic regulators diff...

... genes like glp-1 , skn-1 and pop-1 . By contrast, later in development, cells that are destined to make the pharynx and that deriv...

... that derive from different cell lineages, coordinate their development to form an integrated, functioning structure. The genes tha...

The C. elegans pharynx: a model for organogenesis : 6. The organ selector gene pha-4
... organ selector gene pha-4 The central regulator of pharynx development is pha-4 , a FoxA transcription factor ( Horner et al., 199...

...al., 2006 ). Conversely, expression of pha-4 throughout the embryo can induce pharyngeal fate in a subset of embryonic blastom...

The C. elegans pharynx: a model for organogenesis : 8. Downstream of PHA-4: patterning the pharynx primordium
...latory network that governs the latter stages of pharyngeal development ( Table 2 ). Table 2. Algorithms. Tools for genomic approac...

The C. elegans pharynx: a model for organogenesis : 11. Transcriptional strategies for organogenesis
... pharynx vs. the midgut, two very different organs. Pharynx development depends on PHA-4 , which functions at multiple stages of development and in all pharyngeal cell types. To achieve diversity, pha...

...f one of these GATA factors, end-1 , can convert the entire embryo into midgut ( Zhu et al., 1998 ) and another GATA factor, e...

The C. elegans pharynx: a model for organogenesis : 12. Morphogenesis
...f cells bordering the nascent midgut in the interior of the embryo ( Portereiko and Mango, 2001 ; Sulston et al., 1983 ). The ...

...rane Pha morphology Muriel et al., 2005 ham-2 Zinc finger Z Embryo Nuclear Pha attachment Baum et al., 1999 inx-3 Innexin Z Br...

...d et al., 2006 ). By time-lapse videomicroscopy, pharyngeal development proceeds normally to the 1.5 fold stage when pharyngeal cel...

...-6 suggesting that the regulatory circuit that controls eye development in other animals may have adopted a new function for anteri...

The C. elegans pharynx: a model for organogenesis : 14. Is the pharynx a heart?
.... Second, the involvement of Nkx2.5 proteins for pharyngeal development may be misleading. In other organisms, Nkx2.5 factors are r...

... organisms, Nkx2.5 factors are required for visceral muscle development as well as heart formation (e.g., tinman in Drosophila ( Bo...

...cytes in zebrafish, can nevertheless rescue visceral muscle development in Drosophila Nkx/tinman mutants ( Park et al., 1998 ). Thi...

The C. elegans pharynx: a model for organogenesis : 15. Conclusion
... to visualize individual cells during organogenesis and the development of powerful tools (genomics, forward and reverse genetics, ...

The C. elegans pharynx: a model for organogenesis : 16. Acknowledgements
...all my lab, current and past, for discussions on pharyngeal development. S.E.M. is supported by R01 DK070184 and R01 GM056264 from ...


The C. elegans intestine [HTML]
The C. elegans intestine
...Department of Biochemistry and Molecular Biology, Genes and Development Research Group, University of Calgary, Calgary, Alberta T2N...

...lly, the transcription factor network controlling intestine development and function. ...

The C. elegans intestine : 1. Introduction
...in C. elegans , beginning with the intestinal cell lineage, development and morphogenesis, and then proceeding to the cell biology ...

The C. elegans intestine : 2. The intestinal cell lineage in time and space
...ing from a single cell, the E blastomere, of the eight-cell embryo ( Deppe et al., 1978 ; Sulston et al., 1983 ). The customar...

...ision of the E blastomere takes place on the surface of the embryo and produces an anterior daughter Ea and a posterior daught...

...aughter Ep (see Figure 2 ). Ea and Ep then migrate into the embryo during gastrulation (see Gastrulation in C. elegans ). The ...

The C. elegans intestine : 3. Intestinal morphogenesis and patterning
... and int4, undergo a clockwise rotation (as seen facing the embryo) of approximately 90° ( Sulston et al., 1983 ); this ro...

...IN-12 /Notch pathway (see Notch signaling in the C. elegans embryo ). When the intestine primordium has four cells, the LIN-12...

...t of Notch signaling (see Notch signaling in the C. elegans embryo ; Neves and Priess, 2005 ). The REF-1 effect is ascribed to...

... lineages and indeed the protein is widespread in the early embryo. Finally, we recall the observation that expression of bioc...

The C. elegans intestine : 4. Structure of an intestinal cell
...ical in establishing the fundamental asymmetry of the early embryo (see Gastrulation in C. elegans ), PKC-3 apical localizatio...

...nt phosphatidylserine binding protein) is widespread in the embryo but after hatching, most GEM-4 is located in the intestinal...

... is a gap junction channel protein, widely expressed in the embryo. At the comma stage, INX-3 becomes strongly (if transiently...

...;gut granules” ( Kostich et al., 2000 ). In the early embryo, LMP-1 protein is more widespread and is associated with th...

The C. elegans intestine : 5. Function: towards a molecular physiology of the intestine
...ding genes, is expressed exclusively in the intestine, from embryo to adult, and is induced at the transcriptional level by ex...

...olk into the intestine primordium from the remainder of the embryo ( Bossinger and Schierenberg, 1996 ; Yu et al., 2006 ). Onc...

...for its role in specification of EMS cell fate in the early embryo but SKN-1 has long been suspected of having an additional r...

The C. elegans intestine : 6. Transcriptional control in the intestine
...determining the fate of the EMS blastomere of the four cell embryo (see Figure 2 ; Bowerman et al., 1992 ; Bowerman et al., 19...

...ELT-2 probably contributes to this early phase of intestine development because it is sufficient (though not necessary) for express...

...ion of intestine genes during this early phase of intestine development. The third phase (“Period of ELT-2 Dominance”) ...

...neither factor seems to have a major influence on intestine development, i.e., animals in which the elt-4 gene ( Fukushige et al., ...

The C. elegans intestine : 7. Future directions in the intestine
...Even if the main transcription factors regulating intestine development have been identified, how do they all work together? Will u...


Evolution of development in nematodes related to C. elegans [HTML]
Evolution of development in nematodes related to C. elegans
.... Nematodes of clade IV 3. Developmental systems 3.1. Gonad development 3.2. Vulva development 3.3. Male tail and body size evolution 4. Conclusion 5. Ack...

...es, providing a detailed picture of evolutionary changes in development. In particular, vulva development has been studied in great detail and substantial difference...

...evoldevnematode.html 10.1895/wormbook.1.46.1 Evolution of development in nematodes related to C. elegans Download PDF version Evo...

...des related to C. elegans Download PDF version Evolution of development in nematodes related to C. elegans * Ralf J. Sommer § ...

Evolution of development in nematodes related to C. elegans : 1. Introduction
...pmental processes in more detail. These are vulva and gonad development in hermaphrodites/females, male tail development and body size. Embryonic development and sex determination are covered separately, see Embryolog...

...ariations in a phylogenetic context. For certain aspects of development, such as embryonic, gonad, vulva and male tail development, representatives of up to five nematode families have been ...

...olved? How representative are certain aspects of C. elegans development for other nematodes? What are the directions of evolutionar...

...de, providing a detailed picture of evolutionary changes in development that is unique in the animal kingdom. The following chapter...

Evolution of development in nematodes related to C. elegans : 3. Developmental systems
...en space restrictions, I will focus my description on gonad development, vulva formation, male tail development and body size. I apologize to those colleagues whose work I...

...va formation is one of the best studied processes in animal development (see Vulval development ). Initial genetic and molecular analyses revealed the invo...

...rities to oncogenesis initiated a boost in studies on vulva development. Vulva development occurs in the late L3 and L4 stage. All cell divisions take...

...ald, 2005 ). Over the years, the detailed analysis of vulva development became a paradigm for signaling processes in animal development and thus, provides a platform for studying evolutionary dev...


Roles of chromatin factors in C. elegans development [HTML]
Roles of chromatin factors in C. elegans development
...5/wormbook.1.139.1 Roles of chromatin factors in C. elegans development Download PDF version Roles of chromatin factors in C. elega...

...wnload PDF version Roles of chromatin factors in C. elegans development * Mingxue Cui and Min Han § Howard Hughes Medical Inst...

...romatin factors regulate hox gene expression to control the development of rays in the male tail 3.5. Chromatin factors and silenci...

...tion of the chromatin structure is essential for the proper development of organisms. C. elegans is a powerful organism for explori...

Roles of chromatin factors in C. elegans development : 2. Chromatin factors in C. elegans
..., germline-soma distinction pyp-1 NURF38 ortholog Embryonic development rba-1 NURF55 ortholog Embryonic development, RNAi SWR1/SRCAP complex (ATP-dependent chromatin-remodelin...

... Histone H3K27 methylation, Germline maintenance, Male tail development, RNAi sop-2 SAM domain Male tail development Trithorax group lin-49 PHD finger, bromodomain Hindgut and ...

...complexes have been found to play essential roles in vulval development, germline development, germline-soma distinction, repetitive transgene silencing ...

...ne silencing in both germline and soma, RNAi, somatic gonad development, and larval development ( Andersen et al., 2006 ; Bender et al., 2004a ; Cardoso et...

Roles of chromatin factors in C. elegans development : 3. Chromatin factors regulate a number of cellular and developmental processes
...romatin factors regulate hox gene expression to control the development of rays in the male tail Male tail ray development provides an ideal model system for investigating how hox ge...

...cesses Chromatin factors play an essential role in germline development, which is described in detail in the Germline chromatin cha...

...the role of chromatin factors in several aspects during the development of somatic tissues. 3.1. Several chromatin factors neg...

...l signaling and developmental pattern formation (see Vulval development ). The differentiation of vulval cell types from multiple c...

Roles of chromatin factors in C. elegans development : 4. Summary and future directions
...tin factors also play a role in genome stability, embryonic development, foregut development, etc. ( Pothof et al., 2003 ; Updike and Mango, 2006 ). An ...


Embryological variation during nematode development [HTML]
Embryological variation during nematode development
...895/wormbook.1.55.1 Embryological variation during nematode development Download PDF version Embryological variation during nematod...

...ownload PDF version Embryological variation during nematode development * Einhard Schierenberg § Zoological Institute, Univers...

... studied nematodes leading to the assumption that embryonic development shows little variation within the phylum Nematoda. However,...

Embryological variation during nematode development : 1. Overview
...ail (see Asymmetric cell division and axis formation in the embryo ). As the early cell lineages and arrangement of blastomere...

... implicit assumption has been that the pattern of embryonic development is very similar among nematodes. However, as we know now, t...

Embryological variation during nematode development : 2. Reproduction and diploidy
...985 ) indicates that a diploid status is crucial for normal development. How diploidy is achieved in embryos of three parthenogenet...

...nce visualizing DAPI-stained nuclei. a, C. elegans , 1-cell embryo prior to fusion of pronuclei; two PBs have been extruded at...

...D. coronatus , 2-cell stage, one PB present; insert: 2-cell embryo after division of the single PB; c, A. nanus , 3-cell stage...

Embryological variation during nematode development : 5. Cell lineages
...Cell lineages The impressively detailed studies on cellular development in the Ascaris (clade III) embryo more than a hundred years ago ( Muller, 1903 ) strongly sug...

Embryological variation during nematode development : 7. Cell-cell communication and cell specification
...ell (see Asymmetric cell division and axis formation in the embryo ). Only few data are available on experimental analysis of ...

...somatic lineage have been detected (see sections 4 and 5 ), development of partial embryos and cell marking experiments ( Voronov a...

...ans (see Asymmetric cell division and axis formation in the embryo ), is not required for proper specification of the gut prec...

...lated A. nanus embryos.  Starting from an intact early embryo (top) with the sequentially generated somatic cells AB , EM...

Embryological variation during nematode development : 8. Gastrulation
...on Gastrulation as seen in C. elegans leads to a triblastic embryo with only a rudimentary blastocoel present. It can be subdi...

...ype of gastrulation and other developmental parameters like embryo size, cell numbers or developmental tempo ( Schierenberg, 2...

...he E cell (gut founder) have moved into the interior of the embryo after the 24-cell stage, only rudimentary blastocoel presen...

...t. b, Plectus sp. (ES 601). The E cell has moved inside the embryo after the 12-cell stage. c, Enoplus brevis , 16-cell stage,...

Embryological variation during nematode development : 9. Concluding remarks
...more variable than later phases. One argument has been that development is modular and integration of the emerging modules increase...

...les increases over time, putting fewer constraints on early development ( Raff, 1996 ). This seems reasonable for organisms where c...

...nematodes like Enoplids. However, in the C. elegans type of development, essential decisions requiring intercellular communication ...


Autophagy in C. elegans [HTML]
Autophagy in C. elegans
...gy in C. elegans 3.1. Survival during starvation 3.2. Dauer development 3.3. Reproductive development 3.4. Lifespan extension 3.5. Programmed cell death 3.6. Nec...

...utophagy in an intact multicellular organism that undergoes development to produce different cell types. This review summarizes imp...

Autophagy in C. elegans : 1. Introduction to evolutionarily conserved features of autophagy
... the autophagy pathway and its biological roles in metazoan development and biology. Figure 1: Schematic diagram of the steps of au...

...ther autophagy gene mutants, with respect to abnormal dauer development and mislocalization of the autophagy marker, GFP-LGG-1 ( Me...

...tion, causes developmental arrest at the L3 stage of larval development, with gonadal and intestinal degeneration and increased hyp...

...on 3.4 ). In C. elegans , two other pathways regulate dauer development and thus could potentially regulate autophagy: the guanylyl...

Autophagy in C. elegans : 2. Tools to study autophagy in C. elegans
...ls Our understanding of the role of autophagy in C. elegans development is emerging from studies using either chromosomal mutations...

...s done for bec-1 to uncover its role in longevity and dauer development ( Meléndez et al., 2003 ), as well as RNAi of unc-51...

... , and atg-18 to uncover the role of autophagy during dauer development ( Meléndez et al., 2003 ). RNAi is also accomplished...

...nd ATG16 have still to be tested for a role in autophagy or development ( Table 1 ). C. elegans orthologs to the recently reported ...

Autophagy in C. elegans : 3. Functions of autophagy in C. elegans
...with overactive, non-physiological levels). 3.2. Dauer development The first evidence for a role of autophagy in C. elegans development was provided by studies in daf-2(e1370) mutant dauer animal...

...d recovery defects of pcm-1 mutants. 3.3. Reproductive development The role of autophagy in early development in C. elegans is not confined to developmental changes that...

...assada and Russell, 1975 ; Riddle and Albert, 1997 ). Dauer development is regulated in a temperature-sensitive manner by different...

...If conditions improve, dauer larvae can resume reproductive development, reach the adult stage, and have a normal lifespan. Althoug...

Autophagy in C. elegans : 4. Conclusion
... autophagy is involved in cell survival, cell death, normal development, dauer development, negative control of cell growth, longevity, clearance of t...

...the role of autophagy as an essential process in C. elegans development has advanced greatly in the past few years. We have learned...


Heterotrimeric G proteins in C. elegans [HTML]
Heterotrimeric G proteins in C. elegans
...d EGL-30 are involved in diverse and fundamental aspects of development and behavior. GOA-1 acts redundantly with GPA-16 in positio...

... from the first larval stage. In addition to their roles in development and behaviors such as egg laying and locomotion, the EGL-30...

Heterotrimeric G proteins in C. elegans : 1. Introduction
...l tip cells Early embryos Cell membranes and centrosomes in embryo, neuronal processes in adults Mendel et al., 1995 ; S&eacut...

...maternally for proper spindle positioning in the developing embryo ( Gotta and Ahringer, 2001 ; Tsou et al., 2003 ). GSA-1 and...

...ronal migration, spicule protraction, and may affect vulval development ( Trent et al., 1983 ; Brundage et al., 1996 ; Lackner et a...

...esponse to volatile anesthetics, and may also affect vulval development ( Mendel et al., 1995 ; Ségalat et al., 1995 ; Frase...

Heterotrimeric G proteins in C. elegans : 2. Gαs
...xpressed in neurons and muscles from the embryonic stage of development onward. It may also be expressed in intestinal and some epi...

Heterotrimeric G proteins in C. elegans : 3. Gαq
...yngeal, and defecation muscles and arrest throughout larval development ( Brundage et al. 1996 ). Less severe alleles cause a varie...

...aptic muscle cells to neurons, and for its effect on vulval development ( Lackner et al., 1999 ; Doi and Iwasaki, 2002 ; Moghal et ...

Heterotrimeric G proteins in C. elegans : 5. Gαo
...ns have also revealed roles for GOA-1 in egg laying, vulval development, embryonic viability, locomotion, fertility, and the regula...

... all cells (up to at least the 20-cell stage). In the early embryo, GOA-1 is also localized to the cell cortex and to regions ...

...units causes multiple defects in cell cleavage in the early embryo. During the first mitotic cleavage, nuclear rotation does n...

...lei and few cells ( Srinivasan et al., 2003 ). In the early embryo, GOA-1 and GPA-16 partner with the G β subunit GPB-1 a...

Heterotrimeric G proteins in C. elegans : 6. Gα12
...the G α q and G α 13 pathways to affect embryonic development ( Gu et al., 2002 ). Vertebrate G α 12 is capable of t...

...sis of CeRhoGEF indicates that it plays a role in embryonic development ( Yau et al., 2003 ). ...

Heterotrimeric G proteins in C. elegans : 7. GPAs
...a-3 (QL) transgenics may be due to defects in morphological development of the amphid sensory neurons since these animals are defec...

...onse ( Zwaal et al., 1997 ). In addition to promoting dauer development, pheromone also inhibits recovery of dauer larvae in the pr...

... daf-1 and daf-8 ( Zwaal et al., 1997 ). 7.4.3. Vulval Development GPA-5 acts downstream of the GPCR SRA-13 to negatively regu...

... downstream of the GPCR SRA-13 to negatively regulate vulva development by negatively regulating RAS/MAPK signaling. sra-13 (lf) mu...


Specification of the germ line [HTML]
Specification of the germ line
...important themes have emerged in specifying and guiding the development of the nascent germ line. At early stages, the germline bla...

Specification of the germ line : 1. Overview
...sis (see Asymmetric cell division and axis formation in the embryo ). During early development of the embryo ( Figure 1 ), through a series of asymmetric partitioning e...

...o the germline blastomeres, is crucial for their subsequent development. This chapter discusses the molecular nature and likely rol...

Specification of the germ line : 2.
...er protein with a dynamic localization pattern in the early embryo ( Mello et al., 1996 ; Figure 1 ). PIE-1 accumulates in the...

...96 ; Figure 1 ). PIE-1 accumulates in the oocyte and 1-cell embryo and during each of the initial four divisions becomes enric...

...03 ; see Asymmetric cell division and axis formation in the embryo ). The latter is accomplished by an elongin C/CUL-2 E3 ubiq...

.... pie-1 gene products are maternally provided to the 1-cell embryo along with numerous gene products that encode somatic diffe...

Specification of the germ line : 3. The MES proteins and regulation of chromatin
...e factors that are maternally supplied and that function in development of the germ line in offspring. Indeed, mes/mes mothers prod...

...offspring whose germ line degenerates midway through larval development ( Capowski et al., 1991 ; Paulsen et al., 1995 ). Interesti...

...re maintained in a silent state throughout most of germline development. The concentration of the repressive H3K27me3 mark on the X...

Specification of the germ line : 4. P granules and regulation of RNA
...anules, called P granules, are indeed required for germline development (e.g., Kawasaki et al., 1998 ; Kuznicki et al., 2000 ). The...

... they made of, and what roles do they serve during germline development? P granules are maternally loaded into the oocyte and progr...

...04 ; see Asymmetric cell division and axis formation in the embryo ): 1) posterior movement with bulk cytoplasmic flow, 2) hit...

... is their primary residence throughout the rest of germline development. When considering what roles P granules are likely to serve...

Specification of the germ line : 5. mep-1 and avoiding germline specification
...ermline patterns of gene expression. All cells of the early embryo inherit that germline chromatin state from the oocyte and s...

...e to persist in somatic cells, leading to defective somatic development. Concomitant loss of the MES's would eliminate the germline...

... make MEP-1 dispensable in somatic cells, restoring somatic development. Loss of pie-1 would allow MEP-1 to remodel the germline ch...

... state in the germline cells, leading to defective germline development. Tests of this model will reveal whether MEP-1 in the soma ...

Specification of the germ line : 6. Summary and future directions
...ental germ line and transmission of that state to the early embryo (via the MES system); preservation of that state specifical...


Germline chromatin [HTML]
Germline chromatin
... to an apparent sperm-imprinted X inactivation in the early embryo. Other potential roles for germline-specific modes of chrom...

Germline chromatin : 1. General considerations
...ished in early embryogenesis and then maintained throughout development and in the adult gonad. Key processes for establishment and...

...c germ line, unlike the primordial germ cells (PGCs) in the embryo, is both proliferative and highly transcriptionally engaged...

Germline chromatin : 3. Embryonic germline chromatin
...tiation, phospho-Ser5, is present in all cells of the early embryo, whereas a phospho-epitope corresponding to transcriptional...

...nt chromatin structure is present in all cells of the early embryo, but absent from Z2 and Z3 .  Fixed, whole-mount embry...

...d in these cells, and these are detected at a later time in development (e.g., pgl-1 and nos-2 mRNAs; Kawasaki et al., 1998 ; Subra...

...ux, 1999 ). These cells are mitotically quiescent until the embryo hatches, which may indicate continued transcriptional and/o...

Germline chromatin : 4. Post-embryonic germline chromatin
...al → transition Pachytene Diplotene Diakinesis 1-cell embryo H4acetylK8, K16 Absent Absent Present (low) Present Present...

...#8594; transition Pachytene 1°/2° Spermatids 1-cell embryo H4acetylK8, K16 Absent Absent Absent Absent Present H3diace...

...al → transition Pachytene Diplotene Diakinesis 1-cell embryo H4acetylK8, K16 Present Present Present Present Present H3d...

...#8594; transition Pachytene 1°/2° Spermatids 1-cell embryo H4acetylK8, K16 Present Present Present (low) Absent Presen...

Germline chromatin : 5. Completing the cycle: the zygote
...n the oocyte occur after fertilization, and thus the 1-cell embryo is essentially still very much a germ cell in character wit...

...ke longer (i.e., more cell divisions) to be replaced in the embryo. One obvious difference between these Xp's is that the Xp f...

...vel may subsequently take more rounds of replication in the embryo for effective S-phase coupled histone replacement of the im...


Transcription mechanisms [HTML]
Transcription mechanisms
.... elegans 3. Global transcriptional repression in the early embryo 4. Analyses of GTF functions in C. elegans 5. The many face...

...fferentiation and functions of eukaryotic cells, and to the development of complex organisms. mRNAs are synthesized by the coordina...

...transcription co-factors plays a central role in C. elegans development, and in which C. elegans studies have provided new insights...

Transcription mechanisms : 2. Tools for studying transcription in C. elegans
... ( Section 3 ). It has therefore been possible in the early embryo to study functions of GTFs that are otherwise essential for...

...vel transcription begins during embryogenesis and germ cell development, and to track mRNA localization. A major advance for studyi...

...jor advance for studying C. elegans gene regulation was the development of techniques for visualizing the presence of individual mR...

...a transcription factor, in this case the regulator of Dauer development and longevity DAF-16 ( Oh et al., 2006 ). As a greater numb...

Transcription mechanisms : 3. Global transcriptional repression in the early embryo
...21.1 3. Global transcriptional repression in the early embryo Across diverse species, mRNA transcription is globally shut...

...wn during oogenesis then reactivated early during embryonic development ( Davidson, 1986 ). In C. elegans , this shutdown occurs as...

...ence is then maintained until the 4-cell stage of embryonic development ( Seydoux and Dunn, 1997 ). It is not understood in any spe...

...ional silencing is established, or is later relieved in the embryo. One rationale for having this period of transcription sile...

Transcription mechanisms : 4. Analyses of GTF functions in C. elegans
...ription factors can be investigated in the early C. elegans embryo because of the presence of maternal gene products. For exam...

...n and elongation ( Takagi et al., 2003 ; Figure 1B ). Other embryo studies have studied the core promoter recognition factor T...

...hromatin organization during early stages in the C. elegans embryo, when gene expression patterns are changing rapidly ( Baugh...

Transcription mechanisms : 5. The many faces of Mediator
...T-12 ) acts antagonistically to Ras signaling during vulval development ( Moghal and Sternberg, 2003 ), and that DPY-22 ( MDT-12 ) ...

Transcription mechanisms : 6. Chromatin regulation, an emerging field in C. elegans
...y critical roles in C. elegans transcription regulation and development. The transcription regulators CBP-1 and HDA-1 provide an in...

... Figure 1A ; Chan and La Thangue, 2001 ). In the C. elegans embryo, CBP-1 is required for differentiation of essentially all c...

...ription factor that initiates endoderm specification in the embryo ( Calvo et al., 2001 ; Shi and Mello, 1998 ). Like CBP-1 , ...

...been implicated in additional functions that include vulval development and suppression of inappropriate germline gene expression (...

Transcription mechanisms : 7. Conclusions
...10.1895/wormbook.1.121.1 7. Conclusions Development of an organism depends upon finely-tuned and accurate contr...


Hermaphrodite cell-fate specification [HTML]
Hermaphrodite cell-fate specification : 1. Introduction
... these sexually dimorphic structures are made during larval development by the sex-specific divisions of just 16 blast cells ( Figu...

...ure 4 ). Figure 2. Hermaphrodite and male P lineages.  Development of the ventral nervous systems. Dotted lines indicate the t...

Hermaphrodite cell-fate specification : 2. Basic body plan and role of Hox genes
...( Streit et al., 2002 ), particularly in the case of vulval development (see Vulva development ; Eisenmann et al., 1998 ). As in other systems, the Hox ge...

...sms (see Asymmetric cell division and axis formation in the embryo , Gastrulation in C. elegans and Epidermal morphogenesis )....

...-posterior sister cells in many divisions during C. elegans development ( Herman, 2002 ; Herman and Wu, 2004 ; Lin et al., 1998 ). ...

...in-39 , mab-5 and egl-5 are only required for postembryonic development ( Kenyon et al., 1997 ). Furthermore, it appears that unlik...

Hermaphrodite cell-fate specification : 3. HSN neuron
...ll death plays a large role in shaping the body plan during development, it does not play a large role in shaping the differences b...

.... The HSNs are not needed in males and die during embryonic development. By contrast, the four CEMs are male-specific head neurons ...

...during embryogenesis ( Sulston and Horvitz, 1977 ; see Male development ). HSN death in males is controlled by sex-specific express...

Hermaphrodite cell-fate specification : 4. Epidermal cell fates
...ake-up the vulval precursor cells and participate in vulval development during the third larval stage (see Vulva development chapter for details). In males, P(1–2).p and P(7̵...

... there are many sex-specific specializations, including the development of the hermaphrodite vulva from the ventral hypodermis and ...

... that are derived from only four blastomeres of the 12-cell embryo. The GATA transcription factor ELT-1 appears to be required...

...ically in a similar stem cell-like manner throughout larval development; the anterior daughter fuses with the hypodermal syncytium,...

Hermaphrodite cell-fate specification : 5. Somatic gonad
...he male gonad develops into a single-armed testis (see Male development ). Thus, the SGPs undergo sex-specific cell lineages ( Kimb...

...from animal to animal ( Kimble and Hirsh, 1979 ). Figure 8. Development of the hermaphrodite somatic gonad.  (L1, upper) All c...

...sociated with the germ line and play a critical role in its development and its organization and function in the adult. Reprinted w...

Hermaphrodite cell-fate specification : 7. Conclusions
...ental strategies that differentiate hermaphrodite from male development, some trends are evident. Sex-specific differences in the p...

... go a long way to understanding the control of sex-specific development. Many C. elegans papers on cell lineage or cell fate specif...

Hermaphrodite cell-fate specification : 8. Acknowledgements
...wormbook.1.39.1 8. Acknowledgements Work on C. elegans development in the Herman lab is supported by NIH grant GM56339. This p...


Male development [HTML]
Male development
...maledevelopment.html 10.1895/wormbook.1.33.1 Male development Download PDF version Male development * Scott W. Emmons § Department of Molecular Genetics, ...

...phrodite reproductive structures arise during postembryonic development 2. The major male mating structures form just before the la...

..., and lin-12 signaling specify cell fates in the hindgut 5. Development of the male sensilla 5.1. Rays 5.2. The hook and hook sensi...

...gans male from the hermaphrodite arise during postembryonic development. The major male mating structures, consisting of the blunt ...

Male development: 1. Male and hermaphrodite reproductive structures arise during postembryonic development
...phrodite reproductive structures arise during postembryonic development An L1 larval male just hatched from the egg is not easily d...

...e of the adult male and its postembryonic cell lineages and development were described by Sulston and coworkers ( Sulston et al., 1...

...tarting points for any survey of C. elegans male anatomy or development. In the present chapter, the results of studies on the gene...

...udies on the genetic, cellular, and morphogenetic basis for development of each structure or feature, in so far as they are known, ...

Male development: 4. Multiple pattern formation mechanisms specify cell fates in the tissues that generate the reproductive structures
...ructure has been identified analogous to the anchor cell in development of the hermaphrodite vulva (see Vulval development ; Sulston et al., 1980 ; Sulston and White, 1980 ). In the ...

... inducing tissue (such as the gonad), of male-specific tail development or of any male structure has been identified analogous to t...

...the major hypodermal mating structures during postembryonic development.  The locations of the blast cells in the posterior of...

...vents expression of the Hox gene mab-5 , which promotes ray development ( Hunter et al., 1999 ). There is a directional bias to the...

Male development: 5. Development of the male sensilla
...10.1895/wormbook.1.33.1 5. Development of the male sensilla 5.1. Rays 5.1.1. Ray extensi...

...uring embryogenesis and remains ON throughout postembryonic development except in the ray 3, 5, and 6 branches. (C) EGL-5 expressio...

...n.p cells that generate the hermaphrodite vulva (see Vulval development ), the fates of P10.p and P11.p are established by cell sig...

...s postembryonically During the second half of postembryonic development, the posterior sensory structure common to both sexes, the ...

Male development: 7. Male-specific neuronal circuitry develops during L4
...neurons in the male tail that are born during postembryonic development are the last neurons in the nervous system to differentiate...

Male development: 9. The cell lineage that gives rise to the asymmetric male gonad is a variation on that of the hermaphrodite
...metric, that of the male gonad is one-armed and asymmetric. Development of asymmetry in the male gonad appears already after the se...

...lin-12 and the Wnt receptor lin-17 , are required for gonad development in both sexes. One transcription factor that appears to pla...


LIN-12/Notch signaling in C. elegans [HTML]
LIN-12/Notch signaling in C. elegans
... /Notch family mediate cell-cell interactions during animal development, and aberrations in LIN-12 /Notch signaling have been impli...

LIN-12/Notch signaling in C. elegans : 1. Introduction
...dulated. New roles for LIN-12 /Notch signaling in mammalian development and disease have been identified at a prodigious rate ( Gri...

LIN-12/Notch signaling in C. elegans : 2. Genetic identification and characterization of LIN-12 and GLP-1
...nesis, vulval precursor cell fate specification (see Vulval development ), and sex mesoblast specification (see Male development ; Greenwald et al., 1983 ). lin-12 also is required for ...

..., 1994 ), and at least four other interactions in the early embryo (see also Notch signaling in the C. elegans embryo ). Double mutants lacking both lin-12 and glp-1 activity re...

...n a genetic screen for mutations that have defective vulval development ( Greenwald et al.,1983 ; Ferguson and Horvitz,1985 ). The ...

...d Kimble, 1991 ; see also Notch signaling in the C. elegans embryo ). The functional redundancy inferred from genetic analysis...

LIN-12/Notch signaling in C. elegans : 4. Identification and
...specification ( Chen and Greenwald, 2004b ; see also Vulval development ). 4.2. Site 2 (extracellular) cleavage sup-17 was ide...

...nct roles for transcriptional repression by CSL proteins in development (e.g., Morel and Schweisguth, 2000 ). Association with the ...

LIN-12/Notch signaling in C. elegans : 5. Identification of modulators of
...rdon et al., 2000 ). ego-1 plays multiple roles in germline development, and encodes an RNA-directed RNA polymerase ( Smardon et al...

...s are resistant to RNAi, suggesting a link between germline development and cellular processes involved in post-transcriptional sil...

... proteins may be part of the switch from mitotic to meiotic development (E. Maine, personal communication). The gene atx-2 was also...

LIN-12/Notch signaling in C. elegans : 6. Lateral specification and induction mediated by LIN-12 and GLP-1
...lsewhere in WormBook (see Notch signaling in the C. elegans embryo , Vulval development , Male development , Hermaphrodite cell-fate specification , and Germline prol...

...ate many different cell-cell interactions during C. elegans development. Some of these interactions are considered elsewhere in Wor...

... be unique mediators of lateral specification during animal development. The archetypal example of lateral specification in C. eleg...

...eceptor systems, also mediate inductive interactions during development. In these cases, receptor activity is essentially regulated...

LIN-12/Notch signaling in C. elegans : 7. Future prospects
... regulatory circuits (see Notch signaling in the C. elegans embryo ). Modulation is likely to involve different mechanisms for...


Small GTPases [HTML]
Small GTPases
.... Rho 2.5. Cdc42 2.6. Rac 2.7. Overlapping roles of Racs in development 2.8. Modularity of Rac signaling 2.9. Rab family 2.10. Ran-...

...rphogenesis and microtubule organization and possibly cilia development. Functions for many of the small GTPases remain to be disco...

...egans will elucidate the roles of these molecules in animal development. ...

Small GTPases : 1. Ras-superfamily GTPases in C. elegans
...about the roles of Ras and Rho family GTPases in C. elegans development, while many other GTPase families (e.g., Rab) remain unchar...

...haracterized. Key to understanding small GTPase function in development will be to determine the differential roles of GTPases and ...

Small GTPases : 2. Ras/Ral/Rap family GTPases
...GTPases. The roles of many of these molecules in C. elegans development remain mysterious. The K-Ras homolog and the Rap subfamily ...

... ). A detailed description of let-60 function in C. elegans development can be found in RTKRas/MAP kinase signaling . 2.2. Rap...

...in C. elegans . While Rho GTPases each have unique roles in development, an emerging theme in Rho GTPase function is that Rho GTPas...

...00 ; Spencer et al., 2001 ). Embryos that arrested later in development displayed severe defects in tissue morphogenesis. Other mut...

Small GTPases : 3. Summary and future directions
...red to decipher the roles of all Ras-superfamily GTPases in development and to delineate the signaling pathways by which they contr...


Sarcomere assembly in C. elegans muscle [HTML]
Sarcomere assembly in C. elegans muscle : 1. Introduction
...ditis elegans has proven to be a useful system to study the development of muscle ( Waterston, 1988 ; Moerman and Fire, 1997 ). Her...

...ise after the end of gastrulation (at 290 min. of embryonic development; Sulston et al., 1983 ) and are defined by the accumulation...

Sarcomere assembly in C. elegans muscle : 2. Muscle attachments
...ttachments in embryos and adults. (E) A wild-type, 420 min. embryo. PAT-4 ::GFP localizes to body-wall muscle attachments (arr...

... (arrow). The scale bar represents 2 μ m. (F) The same embryo as that shown in (E) double stained with monoclonal antibod...

Sarcomere assembly in C. elegans muscle : 4. Initiating a sarcomere leads to distinct assembly dependence pathways for dense
... in the unc-52 gene block all subsequent steps in sarcomere development, including the next step, which is the polarization of inte...

...LIM proteins are found within the nucleus throughout muscle development and growth ( Figure 6 ). Whether any of these proteins shut...

Sarcomere assembly in C. elegans muscle : 6. Spacing of the components
.... Figure 8. View of a phalloidin stained 1.5 fold wild type embryo.  (A) Whole embryo shown from a left dorsal view. Two bundles of actin filamen...

...cessary for cell motility. During nematode body wall muscle development integrin clusters must also have some flexibility. This can...

...ents. Laser ablation of muscle precursors in the developing embryo leave substantial gaps in the normally continuous muscle qu...

...l just across the basement membrane, but at later stages of development myotactin moves from these nearly longitudinal lines into a...

Sarcomere assembly in C. elegans muscle : 7. Is sarcomere assembly in C. elegans a general model of sarcomere assembly?
...embly in C. elegans have any relevance for mammalian muscle development. For several years those working on myogenesis in vertebrat...

Sarcomere assembly in C. elegans muscle : 8. Summary
...between sarcomere units generated and maintained throughout development? 3. How are actin and myosin microfilaments inserted into t...


Cell-cycle regulation [HTML]
Cell-cycle regulation
...f CDKs 3.5. Protein degradation 4. Cell-Cycle regulation in development 4.1. Cell-cycle variation 4.2. Checkpoint control 4.3. Cell...

...nces Abstract Cell-division control affects many aspects of development. Caenorhabditis elegans cell-cycle genes have been identifi...

...on with growth, differentiation and tissue formation during development. Results from several studies indicate a critical role for ...

Cell-cycle regulation: 1. Overview
...10.1895/wormbook.1.28.1 1. Overview Animal development from a single-cell zygote to fertile adult requires many ro...

... and reduction of corresponding CDK/cyclin activity. During development, variations of this typical somatic division cycle are used...

...e stereotypical pattern of cell divisions during C. elegans development. Cell external signals and cell intrinsic information toget...

...is cell division temporally and spatially controlled during development, and how is progression through the cycle coordinated with ...

Cell-cycle regulation: 3. Regulators of the cell cycle
...andidate null mutant animals arrest cell division during L1 development. Several observations indicate that the post-embryonic prec...

...1 and cyb-3 each are individually required during embryonic development, simultaneous inactivation of these mitotic cyclins causes ...

...re required for progression through G 1 phase during larval development ( Boxem and van den Heuvel, 2001 ; Park and Krause, 1999 )....

...arresting during embryogenesis, during early or late larval development, and as sterile adults. In other metazoans, Cdk2 acts with ...

Cell-cycle regulation: 4. Cell-Cycle regulation in development
...10.1895/wormbook.1.28.1 4. Cell-Cycle regulation in development 4.1. Cell-cycle variation As metazoans go through development, their cells progress through various types of cell cycles....

... and divide asynchronously. Just a few hours into embryonic development, cells in different lineages diverge greatly in cell-cycle ...

...in the intestinal nuclei occurs between 6 and 8 hours of L1 development, approximately 4 hours before nuclear division ( Boxem and ...

...ake place in the intestine and hypodermis during C. elegans development ( Hedgecock and White, 1985 ). Fourteen of the twenty intes...


Neurogenesis in the nematode Caenorhabditis elegans [HTML]
Neurogenesis in the nematode Caenorhabditis elegans
...pter is in WormBook section: > Neurobiology and behavior >> Development View/Add Comments Publication History version 1 latest vers...

...rve out in this review a few general principles of neuronal development in C. elegans . These principles may be conserved across ph...

Neurogenesis in the nematode Caenorhabditis elegans : 1. Introduction
...are 222 neurons (202 somatic, 20 pharyngeal). During larval development a number of neurons are added resulting in a total number o...

Neurogenesis in the nematode Caenorhabditis elegans : 2. Neuronal cell lineages and neuron classification
...lineage diagram reveals some key features of nervous system development ( Figure 2 ). As a quick glance at the diagram shows, C. el...

...ionship. Due to extensive cell migrations in the developing embryo, neurons in direct proximity to one another are also not ne...

... system develops ( Table 1 provides an overview of neuronal development mutants). Below, I highlight a selected few of these mutant...

...d. Other reviews have ventured more into the details of the development of specific nervous system components, such as the male ner...

Neurogenesis in the nematode Caenorhabditis elegans : 3. Genes controlling lineage decisions
... “neuronal ground state” in vertebrate neuronal development ( Hemmati-Brivanlou and Melton, 1997 ). In vertebrates, the...

Neurogenesis in the nematode Caenorhabditis elegans : 4. Genes controlling neuron class specification
...gene regulatory factors that act at late stages of neuronal development, most often during terminal differentiation of a select num...

...-86 or nhr-67 can have distinct roles early and late in the development of a neuronal lineage. 4.3. Other regulatory routines ...

Neurogenesis in the nematode Caenorhabditis elegans : 6. Linking neuronal class specification to lineage
...eus) daughter of virtually every cell division in the early embryo. Work over the past few years has extended this observation...

...ivisions during later stages of embryonic and postembryonic development within and outside the nervous system (reviewed in Mizumoto...

...tive and predictive, as has been shown at various stages of development through the manipulation of the Wnt signaling system that a...

...rent lineages and at different stages of embryonic neuronal development, revealing the ubiquitous employment of the system ( Bertra...

Neurogenesis in the nematode Caenorhabditis elegans : 7. Conclusions and perspectives
...wed here some general features of C. elegans nervous system development and highlighted some emerging principles. The analysis of m...

...but not all, genes that act at different stages of neuronal development (neuron vs. non-neuron, lineage specification, terminal neu...

...et al., 2010 ). The other key concept that requires further development is the Wnt/ POP-1 system. Where does the ligand come from? ...

...e nature of the regulatory states in each neuroblast during development? Systematic mapping of transcription factor expression prof...


Germline proliferation and its control [HTML]
Germline proliferation and its control
... from one primordial germ cell (PGC) set aside in the early embryo to over a thousand cells in the adult. Most germline prolif...

Germline proliferation and its control: 1. Overview
... from one primordial germ cell (PGC) set aside in the early embryo to over a thousand cells in the adult. Early PGCs are incor...

...rm lines.  The wild-type germ line begins in the early embryo with a single primordial germ cell (PGC), which divides onc...

Germline proliferation and its control: 2. Course of germline proliferation
...ine proliferation The germ line is established in the early embryo with birth of P 4 , the first primordial germ cell (PGC). A...

...the first primordial germ cell (PGC). After moving into the embryo during early gastrulation, P 4 divides to generate two PGCs...

...ne proliferation arrests if larvae enter the dauer stage of development and resumes when dauers reenter the normal life cycle. The ...

Germline proliferation and its control: 3. Control of germline proliferation by Notch signaling and the somatic gonad
...p cells (DTCs) control germline proliferation during larval development and control germline mitoses in the adult. Removal of DTCs ...

...ary and sufficient for germline proliferation during larval development and for maintenance of germline mitoses in the adult. The g...

...tch signaling controls germline proliferation during larval development and maintains germline stem cells in the adult ( Crittenden...

...ux and Schedl, 2001 ; see Notch signaling in the C. elegans embryo ). The core components of the Notch signaling pathway inclu...

Germline proliferation and its control: 4. Control of the mitosis/meiosis decision
... all RNA regulators that control multiple steps in germline development. Here, we briefly summarize their molecular identities and ...

Germline proliferation and its control: 5. Other genes affecting germline proliferation
...line proliferation as well as for other aspects of germline development and early embryogenesis (see Specification of the germ line...

...(NOS) proteins are critical for several aspects of germline development: proliferation (this section), sex determination (see Sex-d...

...mals depleted of nos activity. Furthermore, later in larval development, the three nos genes have overlapping functions required fo...


Genetic dissection of developmental pathways [HTML]
Genetic dissection of developmental pathways : 2. Epistasis analysis
...r example, vulval differentiation in C. elegans (see Vulval development ). In wild-type animals, three of six equipotential vulval ...

... of pathways.  (A) Substrate Dependent pathway: vulval development. A series of genes (1, 2 and 3) are necessary to produce a ...

...eries of outcomes (X, Y and Z). lin-26 is necessary for the development of Pn.p cells. lin-39 is necessary to select a subset of Pn...

Genetic dissection of developmental pathways : 3. Epistasis analysis of switch regulation pathways
...would lead to the inability to activate the male program of development, resulting in the hermaphrodite program of development. Removal of both tra-1 and her-1 would make tra-1 unable to...

...hese genes are also responsible for promoting male germline development and negatively regulating female germline development. In the case of a high X:A ratio, inactive fem-1 , fem-2 , ...

...nt at fem-1 , fem-2 , and fem-3 and realizing that germline development is regulated differently than somatic development, the intersex phenotype of animals carrying the tra-1 mutat...

...r pathway. Although tra-1 is downstream of fem-1 in somatic development, it is not so in germline development. Thus, the order of gene action should be determined by epi...


Genetic dissection of developmental pathways [HTML]
Genetic dissection of developmental pathways : 2. Epistasis analysis
...r example, vulval differentiation in C. elegans (see Vulval development ). In wild-type animals, three of six equipotential vulval ...

... of pathways.  (A) Substrate Dependent pathway: vulval development. A series of genes (1, 2 and 3) are necessary to produce a ...

...eries of outcomes (X, Y and Z). lin-26 is necessary for the development of Pn.p cells. lin-39 is necessary to select a subset of Pn...

Genetic dissection of developmental pathways : 3. Epistasis analysis of switch regulation pathways
...would lead to the inability to activate the male program of development, resulting in the hermaphrodite program of development. Removal of both tra-1 and her-1 would make tra-1 unable to...

...hese genes are also responsible for promoting male germline development and negatively regulating female germline development. In the case of a high X:A ratio, inactive fem-1 , fem-2 , ...

...nt at fem-1 , fem-2 , and fem-3 and realizing that germline development is regulated differently than somatic development, the intersex phenotype of animals carrying the tra-1 mutat...

...r pathway. Although tra-1 is downstream of fem-1 in somatic development, it is not so in germline development. Thus, the order of gene action should be determined by epi...


RTK/Ras/MAPK signaling [HTML]
RTK/Ras/MAPK signaling
...APK) signaling pathways are used repeatedly during metazoan development to control many different biological processes. In the nema...

RTK/Ras/MAPK signaling : 1. Introduction
...APK) signaling pathways are used repeatedly during metazoan development to control many different biological processes ( Schlessing...

..., the best studied of which is vulval induction (see Vulval development ). Genetic screens based on various let-60 mutant phenotype...

...-dependent processes include vulval, excretory duct and P12 development; B) EGL-17 /EGL-15-dependent processes include sex myoblast...

RTK/Ras/MAPK signaling : 3. Phenotypes of Ras pathway mutants
...ith Notch and Wnt signaling to promote hermaphrodite vulval development (see Vulval development ). let-60 loss-of-function mutants lack a vulva (Vulvaless ...

...es. Because of the widespread roles of Ras signaling during development, mutations affecting the Ras pathway can cause many differe...

... male spicule fates ( Chamberlin and Sternberg, 1994 ; Male development ). Reduced Ras signaling causes spicule defects and prevent...

RTK/Ras/MAPK signaling : 4. Screens used to identify Ras pathway components
... focused on the role of the Ras pathway in promoting vulval development (see Vulval development ), and many known components of the pathway have been ident...

RTK/Ras/MAPK signaling : 8. Regulators of Ras activity
...1 and GAP-2 negatively regulate Ras signaling during vulval development and excretory duct development, respectively ( Hajnal et al., 1997 ; Hayashizaki et al., 1...

RTK/Ras/MAPK signaling : 9. Regulators of the Raf/MEK/ERK kinase cascade
... cascade, but that are not individually required for normal development in most tissues. In general, the roles of these genes can o...

RTK/Ras/MAPK signaling : 10. Targets of MPK-1 ERK, and other factors influencing downstream responses
...o function together, and they have strong effects on vulval development and weaker effects on excretory duct and P12 development ( Nilsson et al., 1998 ; Nilsson et al., 2000 ; Singh and H...

...r, and they have moderate effects on excretory duct and P12 development and weaker effects on vulval development ( Howard and Sundaram, 2002 ). EOR-1 and EOR-2 appear to ac...

...Tan et al., 1998 ), which both promotes and inhibits vulval development, the Hox protein LIN-39 ( Clark et al., 1993 ; Eisenmann et...

...Koh et al., 2004 ; Koh et al., 2002 ), which promote vulval development, and the Hox protein EGL-5 ( Chisholm, 1991 ; Jiang and Ste...

RTK/Ras/MAPK signaling : 11. Interactions between the RTK/Ras/MAPK pathway and other signaling pathways
...enmann et al., 1998 ; Gleason et al., 2002 ). During vulval development (see Vulval development ), the Ras pathway also acts sequentially with a Notch path...

RTK/Ras/MAPK signaling : 12. Conclusions and future prospects
...sponds to the same basic signaling pathway. Although vulval development has been and will continue to be a powerful model for study...


Epithelial junctions and attachments [HTML]
Epithelial junctions and attachments
...deal with epithelial cells in the context of the developing embryo. ...

Epithelial junctions and attachments : 2. Approaches to identify junctional and cytoskeletal components
...et al., 2000 ; Legouis et al., 2003 par-3 PAR-3 (PDZ) Early embryo; int., spermT. Apical Emb; gastrulation def.; Ste RNAi Aono...

... al., 2003 ; Nance and Priess, 2002 par-6 PAR-6 (PDZ) Early embryo; int. Apical Emb; gastrulation def. RNAi Nance et al., 2003...

Epithelial junctions and attachments : 3. Epithelial cytoskeleton
... VAB-10A pattern in adults (a FO component). (E) Mid-staged embryo stained with the MH27 mAb showing the AJM-1 pattern (a CeAJ...

...AR-6 see Asymmetric cell division and axis formation in the embryo ). To fit proteins on a single page, some repeats were omit...

...ic sheath, a structure thought to maintain the shape of the embryo during elongation ( Costa et al., 1997 ; Priess and Hirsh, ...

Epithelial junctions and attachments : 5. Cell adhesion, microfilament anchorage and gate function
...bsp; (A) Flat-mount representation of the dorsal half of an embryo (for a more complete view, see Figure 6 ), with seam cells ...

...essed in other epithelia, it only appears essential for the development of epithelia not lined by a cuticle (Göbel et al., 200...

Epithelial junctions and attachments : 6. Assembly of apical junctions and establishment of epithelial polarity
...ets or tubes ( Nelson, 2003 ; Zegers et al., 2003 ). During development of the epidermis, CeAJ components are assembled in a step-w...

...een difficult to assess owing to their early requirement in development ( Pellettieri and Seydoux, 2002 ). Roles for PAR-3 and PAR-...

Epithelial junctions and attachments : 7. Junction disassembly and cell fusion
... fusion Multiple cell fusion events occur during C. elegans development, which directly affect its morphogenesis and organogenesis....

Epithelial junctions and attachments : 8. Fibrous organelles: junctions with the extracellular matrix and mechanical coupling
...s are not attached to the epidermis and collapse inside the embryo; FOs are strongly disorganized/absent. (B – right) At...


Germline genomics [HTML]
Germline genomics
...aphrodites 2.3. Analysis of germline gene expression during development 3. Chromosome bias for genes with germline-enriched express...

...tify genes with enriched expression in the germ line during development. This knowledge provides a database for further studies tha...

...further studies that focus on gene function during germline development or early embryogenesis. Additionally, a comprehensive overv...

Germline genomics : 1. Regulatory mechanisms of germline gene expression
...? For those wishing to understand these aspects of germline development, one goal is to identify every germline-expressed gene. Wit...

Germline genomics : 2. Gene expression profiling of the germ line
...as well as maternally provided factors necessary for proper development of the early embryo. Combination of the glp-4 (ts) vs wild type comparison and ...

...alysis of mutants that perturb specific aspects of germline development has been very useful in defining sets of genes with germlin...

...004 . 2.3. Analysis of germline gene expression during development In addition to expression profiles of mutants, temporal exp...

...have been probed with RNAs prepared at consecutive times of development from wild type animals. This timecourse provides high tempo...

Germline genomics : 4. Integration of germline functional genomic data
...ion pattern of several thousand genes at multiple stages of development, from embryos to young adults, specifically in hermaphrodit...

...nes required for correct localization of PIE-1 in the early embryo ( Pellettieri et al., 2003 ; see Specification of the germl...

...d by RNAi analysis of candidate genes for roles in germline development ( Hanazawa et al., 2001 ). Additionally, protein partners a...

...netic regulatory networks underlying germline and embryonic development. ...

Germline genomics : 5. Future directions
...4.1 5. Future directions Many questions about germline development and function remain that will benefit from genomic analyses...

...ene expression differ between different stages of germ cell development? Many additional mutants exist that can be used to further ...

...enes will shed significant light on their roles in germline development (see for example Colaiácovo et al., 2003 ). Proteomi...


Programmed cell death [HTML]
Programmed cell death
...rogrammed cell death is an integral component of C. elegans development. Genetic studies in C. elegans have led to the identificati...

Programmed cell death: 1. Introduction
...2 ); 3. programmed cell death that occurs during C. elegans development is determined by the essentially invariant somatic cell lin...

...itz, 2003 ; Sulston, 2003 ). Figure 1. Nomarski image of an embryo with apoptotic cells.  Three cells indicated by arrows...

...rrows underwent programmed cell death in a bean/comma stage embryo and exhibit a refractile, raised-button-like appearance. Th...

...erent types of tissues: during embryonic and post-embryonic development of the soma (referred to as "developmental cell death"; Sul...

Programmed cell death: 2. Killing phase
...ll cells that normally undergo programmed cell death during development ( Conradt and Horvitz, 1998 ; Ellis and Horvitz, 1986 ). Fu...

...lls from undergoing programmed cell death during C. elegans development ( Hengartner et al., 1992 ). Loss-of-function mutations in ...

...e the activation of programmed cell death during C. elegans development ( Horvitz, 2003 ). CED-4 has been shown to physically inter...

... which blocks most, if not all programmed cell death during development, impairs the binding of EGL-1 to CED-9 and EGL-1 -induced r...

Programmed cell death: 3. Specification phase
...cification phase Out of the 1090 cells generated during the development of the soma of a C. elegans hermaphrodite, 131 undergo prog...

...ese cells die during embryonic and 18 during post-embryonic development; Sulston and Horvitz, 1977 ; Sulston et al., 1983 ). If pre...

...ast, most of the 131 cell deaths observed during C. elegans development appear to occur in a cell-autonomous manner; i.e. cells app...

...lators of egl-1 expression have additional functions during development. For example, TRA-1 is the terminal, global regulator of so...

Programmed cell death: 4. Execution phase
... causes delayed appearance of embryonic cell corpses during development and reduced cell deaths in sensitized genetic backgrounds, ...

...itz, 1998a ; Wu et al., 2001 ; Zhou et al., 2001 ), and the development of D-type motorneurons and amphid sensory neurons ( Lundqui...


Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses [HTML]
Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses
... the culture in proportion to their cell type in the mature embryo. Differentiated cells survive well for at least 2 weeks. It...

Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses : 1. Introduction
...ques can be executed against the backdrop of known cellular development, defined anatomy and a sequenced well-annotated genome. The...

...d function. In this chapter we review the history of method development, the protocol, and electrophysiological methods for analyse...

Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses : 2. A long-sought method for isolation and culture of embryonic C. elegans cells
... attachment of the cells to substrates, which precluded the development of a reproducible protocol ( Bloom, 1993 ). However, report...

...the 1-cell stage that they used to study the early steps of development ( Edgar, 1995 ). Embryonic cells were cultured by removing ...

Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses : 3. The protocol
...arrow points to the transparent eggshell that surrounds the embryo. ( B ) Photograph of an egg that has been treated with 2 mg...

...l has been digested by the enzymatic treatment and that the embryo is now in direct contact with the egg buffer. ( C ) Photogr...

...ntact with the egg buffer. ( C ) Photograph of a three-fold embryo released from the eggshell by chitinase treatment. ( D ) Em...

Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses : 4. Cultured cells differentiate and behave in vitro as they do in vivo
...ehave in vitro as they do in vivo Early work on small scale embryo preps first revealed that that cultured embryonic cells beh...

...y cells in proportion to their normal proportion within the embryo. For example, there are 81 muscle cells and 222 neurons in ...

...f cells (4 of ~ 560 cells - 0.7%– in an intact 3 fold embryo are touch neurons) ( Figure 2A ; Suzuki et al., 2003 ; Bian...

Culture of embryonic C. elegans cells for electrophysiological and pharmacological analyses : 5. Electrophysiological techniques applicable to C. elegans cultured cells
..., 2002 ). For example it has been reported that the correct development of the sensory process of AWC neurons strictly depends on t...


C. elegans network biology: a beginning [HTML]
C. elegans network biology: a beginning
...rized cell lineage, and powerful tools available to dissect development, Caenorhabditis elegans, among metazoans, provides an optim...

C. elegans network biology: a beginning : 1. Introduction
...consequences of in vivo perturbations promises to drive the development of increasingly sophisticated models of the topology, funct...

...ome sequence and its annotation laid the foundation for the development of a variety of functional genomic or “omic” ap...

C. elegans network biology: a beginning : 2. Network components: transcriptome and ORFeome
...cess to precise gene structural information has enabled the development of multiple genome-scale resources that can be used for a v...

C. elegans network biology: a beginning : 3. Network connections: interactome
...the context of specific biological processes such as vulval development, protein degradation, germline development, DNA damage response and Dauer formation ( Walhout et al., ...

C. elegans network biology: a beginning : 4. Network perturbations: phenome
... set of comprehensive studies, RNAi phenotypes in the early embryo were analyzed using a systematic approach to explicitly sco...

...tified genome-wide to elicit an RNAi phenotype in the early embryo could be placed into 23 separable phenotypic classes ( S&ou...

...to drive early embryogenesis ( Gunsalus et al., 2005 ). The development of automated image analysis could alleviate what is current...

...features. Automated image analysis pipelines are also under development for analyzing cell lineage patterns ( Yasuda et al., 1999 ;...

C. elegans network biology: a beginning : 5. Network dynamics: from transcriptome to localizome
...and protein products are expressed and localized throughout development, both at a cellular and sub-cellular level. Such a “l...

...1) ; Baugh et al. (2003) ; Robertson et al. (2004) Germline development Reinke et al. (2000) ; Reinke et al. (2004) Meiosis Colaiac...

...ution data on spatiotemporal expression patterns throughout development can be obtained using microscopic techniques. A large-scale...

C. elegans network biology: a beginning : 6. Network modeling: “omic” data integration
...ome-wide phenotypic analysis of genes required in the early embryo, a combined analysis of high-content phenotypic data ( S&ou...

C. elegans network biology: a beginning : 7. Perspectives
...and organismal biology as more efforts are dedicated to the development of resources and the generation of improved interactome, ph...

...y in its infancy, and it will be interesting to see how its development will help bridge network biology with the nearly perfect an...


Transcriptional regulation transformationmicroinjection [HTML]
Transcriptional regulation transformationmicroinjection : 2. Tools to study transcriptional regulation
...ression relied on lacZ reporter genes and were aided by the development of a set of vectors by the Fire lab ( Fire et al., 1990 ). ...

...the DNA binding properties of DAF-12 , a regulator of dauer development and lifespan, to define potential binding sites and gene ta...

...ugh homogeneous tissue for biochemical analysis. The recent development of cell culture techniques (see Methods in Cell biology ), ...

Transcriptional regulation transformationmicroinjection : 4. Simple promoters
...ould genes with such dynamic expression profiles throughout development be regulated in an apparently simple way? The answer is lik...

Transcriptional regulation transformationmicroinjection : 5. Complex promoters
... functionally related to factors controlling cardiac muscle development in other species ( Haun et al., 1998 ). The organ-specific ...

...muscle and all other pharyngeal cell types during embryonic development (see below). CEH-22 is not the only factor functioning with...

...ibit defects in B subelement activity and pharyngeal muscle development and function ( Okkema et al., 1997 ). Thus, other as yet un...

...factors will enhance our understanding of pharyngeal muscle development. Figure 1.  CEH-22 and PHA-4 function in combination t...

Transcriptional regulation transformationmicroinjection : 7. Spatial specificity
...Mango, 2002 ). A major question in understanding pharyngeal development is how does PHA-4 regulate genes expressed in different pha...

... pharyngeal cell types and at different times in pharyngeal development. The function of PHA-4 in cell-type specific differentiatio...

...ion of the pharyngeal muscles. Earlier in pharyngeal muscle development, PHA-4 is also required for the initiation of ceh-22 expres...

Transcriptional regulation transformationmicroinjection : 8. Future
...ll be defined. Understanding how these connections regulate development will add an exciting chapter in the study of the worm and f...


Somatic sex determination [HTML]
Somatic sex determination
...d Zarkower § Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, MN 55455 USA This ch...

Somatic sex determination : 1. A global sex determination pathway controls sexual dimorphism
...For further description of male- and hermaphrodite-specific development, see Male development and Hermaphrodite cell fate specification , respectively. F...

...lso controls sex determination in the germ line (see Vulval development ). However, because hermaphrodites need to generate both oo...

Somatic sex determination : 2. The balance of X-linked and autosomal signal elements controls xol-1
...l-1 The sex determination cascade is initiated in the early embryo by the ratio between the number of X chromosomes and sets o...

Somatic sex determination : 4. HER-1 and TRA-2 provide cell non-autonomous control of sexual fate
...nation. HER-1 is a small secreted protein and promotes male development in a cell non-autonomous manner, by inhibiting the function...

...is is of particular importance in the germ line (see Vulval development ), but also is likely to affect TRA-2 expression in the som...

...95 ; Spence et al., 1990 ). All three are required for male development and, based on genetic epistasis tests, the appear to act be...

Somatic sex determination : 5. High versus low tra-1 activity determines somatic cell sexual fates
...minal regulator tra-1 . Active tra-1 promotes hermaphrodite development and prevents male development ( Hodgkin, 1983 ; Hodgkin, 1987 ). tra-1 expresses two mRNA...

...formed into pseudomales that can be fertile, so robust male development is possible in the absence of tra-1 function. However, thes...

...gonads, indicating that some tra-1 activity is required for development of the male somatic gonad ( Hodgkin, 1987 ; Schedl et al., ...

...ex determination function of tra-1 . tra-1 controls gonadal development in concert with at least two other factors. During embryoge...

Somatic sex determination : 6. TRA-1 links global and tissue-specific sexual regulation
...specific sexual regulation TRA-1 directs sexually dimorphic development at the cellular level, with diverse consequences in differe...

Somatic sex determination : 7. Downstream sexual regulators: the interface between sex determination and sexual differentiation
...e male V rays or hermaphrodite lateral hypodermis (see Male development ), but no TRA-1 target has been identified in this lineage....

Somatic sex determination : 8. Some aspects of sex determination may be evolutionarily conserved
...oding DM domain proteins have been shown to regulate sexual development in mammals ( Raymond et al., 2000 ) and fish ( Matsuda et a...

Somatic sex determination : 9. Open questions
...ation interacts with spatial and temporal regulation during development. The improved availability of reporter genes marking sexual...


C. elegans microRNAs [HTML]
C. elegans microRNAs
...f the C. elegans miRNAs are differentially expressed during development indicating a major role for miRNAs in C. elegans development. Given the remarkable conservation of developmental mechani...

...NAs found in many phyla that control such diverse events as development, metabolism, cell fate and cell death. They have also been ...

... lsy-6 and mir-273 , control left-right asymmetry in neural development, and also target key developmental regulators for repressio...

C. elegans microRNAs: 1. Introduction
...ary Ruvkun's group to direct the later stages of C. elegans development in a similar manner to lin-4 ( Reinhart et al., 2000 ). It ...

... recently discovered miRNAs are temporally regulated during development, while others are spatially expressed. Many are highly cons...

C. elegans microRNAs: 2. C. elegans miRNAs
...study of another miRNA, let-7 , which is expressed later in development. let-7 controls the L4-to-adult transition in C. elegans development by interacting with the 3' UTR of multiple target genes, in...

...king up 22 families) and 30 are temporally regulated during development ( Lim et al., 2003 ). lin-4 and let-7 , the first two known...

...t-7 are temporally regulated and govern temporal aspects of development in C. elegans ( Feinbaum and Ambros, 1999 ; Johnson et al.,...

...normal L1-to-L2 transition during C. elegans post-embryonic development ( Ambros and Horvitz, 1984 ) and encodes at least two small...

C. elegans microRNAs: 4. Conclusions
...rent studies that miRNAs play important roles in C. elegans development; while studies in other organisms show that they are import...


Spermatogenesis [HTML]
Spermatogenesis
...o because a simple, chemically defined medium that supports development has been discovered. Unlike nearly all other C. elegans cel...

Spermatogenesis : 1. Overview
... occurs in males, it also occurs during a stage of germline development in the hermaphrodite prior to the onset of oogenesis (see S...

... is transparent and in vitro because a medium that supports development is available ( Machaca et al., 1996 ; Nelson and Ward, 1980...

...are sperm-specific regulators, and genes affecting germline development more generally are described elsewhere (see Sex determinati...

Spermatogenesis : 2. Wild-type spermatogenesis
...e-derived (blue) sperm will produce a mixture of self (pink embryo) and outcross (blue embryo) progeny. (C) A hermaphrodite after being inseminated by ma...

...10.1895/wormbook.1.85.1 2. Wild-type spermatogenesis Development of sperm in C. elegans males has been described in detail (...

... the primary spermatocyte buds off the rachis and completes development without any requirement for intimate association with other...

... male-derived (blue) sperm will produce only outcross (blue embryo) progeny. As noted by others, male-derived sperm are ~50% l...

Spermatogenesis : 3. Identification of spermatogenesis defective mutants
... spermatozoa that enter the egg and the resulting defective embryo dies. (B) The most common terminal stages of mutants that a...

Spermatogenesis : 5. Mutants that affect sperm meiosis
...ith other PUF proteins to maintain viable germ cells during development ( Subramaniam and Seydoux, 1999 ), but it also plays a part...

Spermatogenesis : 10. Post-fertilization mutants
..., but embryogenesis never begins properly and the defective embryo always dies ( Browning and Strome, 1996 ; Hill et al., 1989...


Nomarski images for learning the anatomy, with tips for mosaic analysis [HTML]
Nomarski images for learning the anatomy, with tips for mosaic analysis
...ohn Yochem § Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis MN 55455 USA This cha...

Nomarski images for learning the anatomy, with tips for mosaic analysis : 1. An overview
...he positions of nuclei and of how these nuclei arise during development can be of great help for studies of Caenorhabditis elegans ...

...to ascertain the effects of mosaicism on subsequent growth, development, or fertility. Although the nuclei of the intestine can be ...

Nomarski images for learning the anatomy, with tips for mosaic analysis : 2. The tutorial
...ed to indicate cells that are produced during postembryonic development; the cells that arise during embryonic development do not have a dot in their cell-lineage designations. Also,...

...ial to be specified as progenitors of the vulva (see Vulval development )]. During normal development, the cells usually divide once, and the daughters join the ...

...to take up a large volume of the region. In hermaphrodites, development of the vulva is one of the most intensely studied aspects o...

...an individual. The P cells are dynamic during postembryonic development. The P9.p nucleus shown in the middle panel is actually par...


Sex determination in the germ line [HTML]
Sex determination in the germ line
...system evolve? 6.5. Is C. elegans a good model for germline development in other animals? 6.6. What technical innovations could rev...

... identity is one of a few basic parameters that specify how development should proceed. Although sex determination has profound eff...

Sex determination in the germ line : 1. Sex determination in germ cells
... with regulatory molecules that will direct early embryonic development, including sex determination, and some of these molecules m...

...re superimposed on this core pathway to allow hermaphrodite development. Below, we briefly describe the core pathway and then the a...

Sex determination in the germ line : 2. The soma acts through HER-1 to regulate the sexual fate of germ cells
...s The her-1 gene acts in XO animals to promote male somatic development and continuous spermatogenesis ( Hodgkin, 1980 ). This her-...

Sex determination in the germ line : 4. Special regulatory modules promote hermaphrodite spermatogenesis
...l., 2004 ). Because GLD-1 has pleiotropic roles in germline development ( Francis et al., 1995 ; Francis et al., 1995 ) it must reg...

...o the nucleus. Since tra-2 requires tra-3 to promote female development, and tra-3 encodes a calpain protease ( Barnes and Hodgkin,...

...and TRA-1 help modulate TRA-1 activity during hermaphrodite development. ...

Sex determination in the germ line : 5. Regulation of fem-3 translation is required for hermaphrodite oogenesis
... analysis showed that GLD-3 plays several roles in germline development, one of which is the promotion of spermatogenesis — i...

...tion is that maternal mog activity is sufficient for normal development. Alternatively, the level of fem-3 transcripts in the soma ...

...ago nashi gene, which regulates several aspects of germline development. In C. elegans , mag-1 (RNAi) causes hermaphrodites to prod...

... oocytes, in addition to numerous other defects in germline development ( Ciosk et al., 2004 ; Maine et al., 2004 ). ...

Sex determination in the germ line : 6. Crucial questions for the future
...resulting only because many genes have pleiotropic roles in development? Or do these dual activities help coordinate the overall development of the germ line? Furthermore, do these data shed any light...

...-2 to fem-3 activity automatically flips during late larval development. 6.2. How is the regulation of sexual fate integrated ...

...nation ). 6.5. Is C. elegans a good model for germline development in other animals? Although we have a solid working model of...

...als. Do any of these genes play conserved roles in germline development, like mab-3 and doublesex do in the soma? 6.6. What te...


Sex determination in the germ line [HTML]
Sex determination in the germ line
...system evolve? 6.5. Is C. elegans a good model for germline development in other animals? 6.6. What technical innovations could rev...

... identity is one of a few basic parameters that specify how development should proceed. Although sex determination has profound eff...

Sex determination in the germ line : 1. Sex determination in germ cells
... with regulatory molecules that will direct early embryonic development, including sex determination, and some of these molecules m...

...re superimposed on this core pathway to allow hermaphrodite development. Below, we briefly describe the core pathway and then the a...

Sex determination in the germ line : 2. The soma acts through HER-1 to regulate the sexual fate of germ cells
...s The her-1 gene acts in XO animals to promote male somatic development and continuous spermatogenesis ( Hodgkin, 1980 ). This her-...

Sex determination in the germ line : 4. Special regulatory modules promote hermaphrodite spermatogenesis
...l., 2004 ). Because GLD-1 has pleiotropic roles in germline development ( Francis et al., 1995 ; Francis et al., 1995 ) it must reg...

...o the nucleus. Since tra-2 requires tra-3 to promote female development, and tra-3 encodes a calpain protease ( Barnes and Hodgkin,...

...and TRA-1 help modulate TRA-1 activity during hermaphrodite development. ...

Sex determination in the germ line : 5. Regulation of fem-3 translation is required for hermaphrodite oogenesis
... analysis showed that GLD-3 plays several roles in germline development, one of which is the promotion of spermatogenesis — i...

...tion is that maternal mog activity is sufficient for normal development. Alternatively, the level of fem-3 transcripts in the soma ...

...ago nashi gene, which regulates several aspects of germline development. In C. elegans , mag-1 (RNAi) causes hermaphrodites to prod...

... oocytes, in addition to numerous other defects in germline development ( Ciosk et al., 2004 ; Maine et al., 2004 ). ...

Sex determination in the germ line : 6. Crucial questions for the future
...resulting only because many genes have pleiotropic roles in development? Or do these dual activities help coordinate the overall development of the germ line? Furthermore, do these data shed any light...

...-2 to fem-3 activity automatically flips during late larval development. 6.2. How is the regulation of sexual fate integrated ...

...nation ). 6.5. Is C. elegans a good model for germline development in other animals? Although we have a solid working model of...

...als. Do any of these genes play conserved roles in germline development, like mab-3 and doublesex do in the soma? 6.6. What te...


Ubiquitin-mediated pathways in C. elegans [HTML]
Ubiquitin-mediated pathways in C. elegans
...f ubiquitin pathway components in the context of organismal development. A combination of forward genetics, reverse genetics, and g...

Ubiquitin-mediated pathways in C. elegans: 1. Overview of ubiquitin conjugation
...d pathways are important for multiple aspects of C. elegans development and cellular physiology. ...

Ubiquitin-mediated pathways in C. elegans: 6. Ubiquitin-protein ligases (E3s)
...on and structure, indicating a critical role in presynaptic development ( Schaefer et al., 2000 ; Zhen et al., 2000 ). RPM-1 contro...

...al., 2000 ; Zhen et al., 2000 ). RPM-1 controls presynaptic development by negatively regulating DLK-1 , which functions as the ini...

...h function in the sex determination pathway to promote male development ( Jager et al., 2004 ). SCF FSN-1 : FSN-1 is an F-box prote...

...red to prevent cytoplasmic extensions/blebbing in the early embryo ( Feng et al., 1999 ). 6) A CUL-2 complex containing the SR...


Essential genes [HTML]
Essential genes
...t our ability to analyze essential genes would benefit from development of new tools for conditional inactivation or activation of ...

Essential genes : 1. Overview
...e of about 8500.) With the sequencing of the genome and the development of methods to target specific genes, it is now within our p...

Essential genes : 2. Types of essential genes
... mutation. Sterility could arise due to defects in germline development, somatic gonad development, oogenesis, spermatogenesis, ovulation or fertilization. ...

...d sterile mutations (steriles). Zygotic lethals prevent the development to adult of individuals homozygous for the mutation. Zygoti...

...e a special class of sterilizing mutations that prevent the development of the progeny of hermaphrodites homozygous for the mutatio...

...es whose expression in the mother is required for embryonic development. Sterile mutations prevent the production of fertilized egg...

Essential genes : 3. Methods for identifying essential genes
...0s provided the first stocked lethal mutants for studies of development. Genes identified in these early screens are named either e...

...work were that many genes are required at multiple times in development and maternal gene expression plays a major role in embryoge...

...alski et al., 1982 ; Rose and Baillie, 1980 ). However, the development of free duplications and crossover-suppressing chromosomal ...

...ermaphrodites that produce only unfertilized eggs and whose embryo production can be rescued by mating to wild-type males (for...

Essential genes : 5. Pleiotropic essential gene phenotypes
...s is also required for cell fate specification in the early embryo. However, this function is masked in animals homozygous for...

...s in cell fate specification at multiple times in the early embryo ( Hutter and Schnabel, 1994 ; Mello et al., 1994 ; Moskowit...


Strongyloides spp. [HTML]
Strongyloides spp. : 5. The life-cycle
...rms; this is the free-living adult generation. This type of development is known as indirect, sexual or heterogonic development. The free-living adults mate and the female lays eggs which...

... that they moult via an L2 into infective L3s. This type of development is known as direct, asexual or homogonic development. Infective L3s that have developed via the direct or indire...

...can undergo autoinfection, that is, repeated generations of development in the same host individual. Autoinfection appears to be un...

...ious pathogen of humans. Autoinfection involves accelerated development by larval progeny of parasitic females such that they devel...

Strongyloides spp. : 7. Controlling the life-cycle
...o which indirect (or heterogonic) and direct (or homogonic) development occur in the free-living phases of the Strongyloides life-c...

...ning developing larvae is held external to the host affects development, such that female larvae preferentially develop into free-l...

...erentially develop into free-living adult females (indirect development) at higher temperatures ( Viney, 1996 ). It is probable tha...

...st from which developing larvae are passed also affects the development of the free-living larvae. This occurs in two ways. Firstly...


The evolution of nematode sex determination: C. elegans as a reference point for comparative biology [HTML]
The evolution of nematode sex determination: C. elegans as a reference point for comparative biology
...is pathway links karyotype to phenotype by coordinating the development of sexually dimorphic tissues. In this article, this pathwa...

The evolution of nematode sex determination: C. elegans as a reference point for comparative biology : 2. Genetic and environmental sex determination in nematodes
... of a limiting host factor may be an important cue for male development ( Petersen, 1985 ). Whether the environmental factors that ...

...onic phase) that is similar to Caenorhabditis . Heterogonic development produces males and females from genetically identical L1 la...

...ska, Lincoln Dept. of Entomology. Commitment to heterogonic development in Strongyloides requires amphid neurons homologous to thos...

The evolution of nematode sex determination: C. elegans as a reference point for comparative biology : 3. Deep conservation and cooption in the evolution of nematode sex determination
... The association of DM family member expression with sexual development in cnidarians ( Miller et al., 2003 ) further suggests that...

...ols many key patterning and cell fate decisions in metazoan development, but is conspicuously absent from C. elegans , as judged by...

...on factor, TRA-1 . Perhaps in the distant past, as nematode development became more mosaic and increasingly dependent upon cell lin...

...neage, hh signaling became dispensable for other aspects of development and was coopted into sex determination. Given the unusually...

The evolution of nematode sex determination: C. elegans as a reference point for comparative biology : 4. Conservation of C. elegans pathway components: C. briggsae as the ruler
...Chen et al., 2001 ) mab-3 regulator of male tail and neuron development (male-promoting) DM domain ( Raymond et al., 1998 ) 67% G G...

The evolution of nematode sex determination: C. elegans as a reference point for comparative biology : 6. Comparative functional studies
... key C. elegans genes. Cb-her-1 is required for normal male development, and its overexpression clearly masculinizes XX C. elegans ...

The evolution of nematode sex determination: C. elegans as a reference point for comparative biology : 7. Mating system evolution in Caenorhabditis
... et al., 2004 ). An independent origin of bisexual germline development may help explain discrepancies in germline gene function th...

... that enables hermaphrodite germ cells to change sex during development ( Figure 3A ). Translational repression of tra-2 by the GLD...

...he “sperm on” portion of hermaphrodite germline development, evidence is mounting that the “sperm off” comp...

...While the Cb-fem genes have conserved roles in male somatic development, none have yet been shown to be required for hermaphrodite ...


TGF-β signaling [HTML]
TGF-β signaling
...F- β superfamily ligands play fundamental roles in the development and physiology of diverse animal species. Genetic and genom...

TGF-β signaling: 2. Dauer pathway
...ter more favorable environments, they exit dauer and resume development, entering the L4 stage. Large-scale genetic screens were co...

...to the TGF- β -related pathway that functions in dauer development as the Dauer TGF- β pathway. See appropriate chapters ...

...way. See appropriate chapters for more information on dauer development and the insulin and cyclic nucleotide signaling pathways. 2...

...that the gene has a role in promoting continuous, non-Dauer development. DAF-7 is expressed in the pair of ASI sensory neurons unde...

TGF-β signaling: 3. Sma/Mab pathway
...from mating successfully. For more information on male tail development and function, see chapters on Male development and Male mating behavior . 3.1. DBL-1: the Sma/Mab pat...

...Mab pathway The Sma/Mab pathway regulates body size and the development of male-specific sensory rays and copulatory spicules ( Sav...

...31 . lin-31 was originally identified by its role in vulval development ( Miller et al., 1993 ), but lin-31 mutant males also have ...

...n shown to affect body size but rather play a role in dauer development ( Kimura et al., 1997 ; Jia et al., 2004 ). Mutants in the ...

TGF-β signaling: 4. UNC-129
...role, a redundant role for unc-129 in male tail sensory ray development has been identified ( Ikegami et al., 2004 ). In this pathw...


Mechanism and regulation of translation in C. elegans [HTML]
Mechanism and regulation of translation in C. elegans
...volve changes in the translation of individual mRNAs during development, in response to physiological changes, or after genetic man...

Mechanism and regulation of translation in C. elegans : 2. The Translational machinery
... cytoplasmic PABP. PAB-1 (Y106G6H.2) is essential for gonad development, and PAB-2 is apparently important for somatic development ( Ciosk et al., 2004 ). PAB-3 ( C17E4.5 ) is more similar t...

...on mutants are viable but have defects in growth and larval development. Yeast eIF2A is not an essential protein, but deletion of C...

...ble Egl Neurons Muscle Egl Specific mRNA translation during development References   Jankowska- Anyszka et al., 1998 ; Keiper ...

...legans eIF5 and eIF1 are essential for growth and embryonic development. There are C. elegans homologs for all 12 eIF3 subunits pre...

Mechanism and regulation of translation in C. elegans : 4. Regulation of Translation
...pek-1 in complementary pathways that are essential for worm development, survival, and ER homeostasis. Furthermore, pek-1 mutants h...

...is tra-2 mRNA, whose translation must be repressed for male development. A possible mechanism of action is suggested by the observa...

... translation in germ cells and posterior cells of the early embryo, and one that inhibits repressor activity to promote transl...

...t inhibits repressor activity to promote translation in the embryo ( Marin and Evans, 2003 ). GLD-1 binds to this repressor el...


Intermediary metabolism [HTML]
Intermediary metabolism
...hydrates 5.2. Glyoxylate cycle 6. Metabolic patterns during development and aging 6.1. Development 6.2. Dauer metabolism 6.3. Aging 7. Influence of external f...

...etabolic patterns and activity change as the worm traverses development and ages, or responds to unfavorable external factors, such...

Intermediary metabolism : 1. Introduction
...e most powerful tools for studying the genetics of metazoan development, neurobiology, aging and many other biological processes. T...

Intermediary metabolism : 6. Metabolic patterns during development and aging
...10.1895/wormbook.1.146.1 6. Metabolic patterns during development and aging 6.1. Development Metabolic rate measured as oxygen consumed or heat produced...

...at glyoxylate cycle mRNA and protein increased during early development, peaked at the L2 stage and declined afterwards. Starvation...

...ion induced glyoxylate cycle expression throughout juvenile development ( Khan and McFadden, 1982 ; Liu et al., 1997 ). However, in...

...n expression in dauers is different from that during normal development ( Liu et al., 1997 ). The ATP profiles reflect these change...

Intermediary metabolism : 7. Influence of external factors on intermediary metabolism
...ted with environmental temperature ( Klass, 1977 ), whereas development and metabolic rate show a positive correlation within an ac...

...peratures. However, the rates of complex processes, such as development and behavior, were shown to be also genetically determined....

...cally determined. Clk mutants, which show slow behavior and development, were unable to adjust their growth and behavioral rates to...


Nuclear hormone receptors in C. elegans [HTML]
Nuclear hormone receptors in C. elegans
...tion 8. Xenobiotic response 9. Metabolic control 10. Neural development 11. Future directions 12. Acknowledgements 13. References A...

... lipophilic hormones, which coordinate metazoan metabolism, development and homeostasis. C. elegans has undergone a remarkable expa...

...al timing, diapause, and life span. They also impact neural development, axon outgrowth and neuronal identity. Finally, they influe...

Nuclear hormone receptors in C. elegans : 3. Comparative phylogeny
..., epidermal differentiation, dauer formation DHR3 Embryonic development, molting, metamorphosis ROR alpha Cerebellar differentiatio...

...rmation Tailless A/P patterning, neurogenesis TLX Forebrain development, neural stem cell maintenance NHR-91 No obvious function DH...

...iation Seven-up Photoreceptor fate COUP TF1 COUP TF2 Neural development FAX-1 Neural differentiation dmFAX-1 Unknown PNR Photorecep...

Nuclear hormone receptors in C. elegans : 5. Sex determination
...5/wormbook.1.64.1 5. Sex determination Acting early in development, SEX-1 regulates C. elegans sex determination and dosage co...

Nuclear hormone receptors in C. elegans : 6. The molt cycle
...sis, epidermal seam cell displacement and blunted male tail development ( Kostrouchova et al., 1998 ; Kostrouchova et al., 2001 ). ...

...nd cell elongation. Adults exhibit aberrant somatic gonadal development, tumorous germlines, and are sterile ( Asahina et al., 2000...

Nuclear hormone receptors in C. elegans : 7. Dauer formation
..., TGF-beta and cGMP pathways to mediate either reproductive development or arrest at the dauer diapause. DAF-12 relatives include V...

...s, and directs expression of genes involved in reproductive development, developmental advance, fat metabolism, and accelerated agi...

...-1 /coregulator specify programs of dauer diapause, delayed development, fat storage, and retarded aging (slow life history traits)...

Nuclear hormone receptors in C. elegans : 10. Neural development
...10.1895/wormbook.1.64.1 10. Neural development Because the C. elegans nervous system is described down to ...

...nematode neurobiology. Several identified NRs affect neural development. UNC-55 , an ortholog of the orphan receptor COUP-TF/Seven-...

Nuclear hormone receptors in C. elegans : 11. Future directions
...ion of lipid metabolic networks, the coordination of neural development, and a detailed exploration of the xenobiotic response come...


Wnt signaling [HTML]
Wnt signaling
... V5 polarity 8.7. Polarity of other tail cells 8.8. Spicule development 9. Conclusion 10. Acknowledgements 11. References Abstract ...

... signaling between cells is a conserved feature of metazoan development. Activation of Wnt signal transduction pathways upon ligand...

Wnt signaling : 1. Introduction
...signal transduction pathways used extensively during animal development, from Hydra to humans ( Cadigan and Nusse, 1997 ; Wodarz an...

...nd Polakis, 2000 ). Wnt signals control multiple aspects of development, including the proliferation, fate specification, polarity,...

Wnt signaling : 8. Other processes utilizing Wnt pathway components
...nbsp; POP-1 asymmetry in AB embryonic lineages In the early embryo, most left/right cell divisions generate daughters with equ...

... inherits more MOM-5::GFP. AB descendants isolated from the embryo and allowed to divide in culture for several divisions also...

...chanistic similarity. 8.3. P7.p polarity During vulval development the vulval precursor cells P5.p and P7.p adopt the 2 ° ...

...l., 2004 ). 8.4. 1 ° lineage of P6.p During vulval development, the VPC P6.p adopts the 1 ° fate and generates eight c...

Wnt signaling : 9. Conclusion
...ith other species, Wnt signaling controls a large number of development processes in C. elegans , and the number is still growing. ...


Mitochondrial genetics [HTML]
Mitochondrial genetics
...chondrial dynamics 6. mtDNA copy number 7. Role of mtDNA in development 8. mtDNA mutations 9. mtDNA and aging 10. Perspectives 11. ...

...ital for Caenorhabditis elegans metabolism, physiology, and development. The C. elegans mitochondrial DNA is typical of animal mito...

Mitochondrial genetics: 3. mtDNA-encoded proteins
...r of genetic screens for defects in motility, reproduction, development, and other functions. Perhaps the nematode can better toler...

Mitochondrial genetics: 4. Mitochondrial DNA inheritance
...A may have a significant impact on evolution and on disease development. ...

Mitochondrial genetics: 6. mtDNA copy number
...legans has been investigated ( Tsang and Lemire, 2002 ). An embryo contains ∼ 25,000 copies of mtDNA and this number rem...

... a somatic component is glp-1 independent and occurs during development from the L3 ( ∼ 25,000 copies) to the L4 stage ( W...

...e majority of organellar genomes are associated with oocyte development and the sperm-associated component is minor. When determine...

...0, respectively. The increases in mtDNA copy numbers during development are suggestive of parallel increases in the numbers of orga...

Mitochondrial genetics: 7. Role of mtDNA in development
...10.1895/wormbook.1.25.1 7. Role of mtDNA in development A functional MRC is essential for viability and larval development past the L3 stage. Mutations in nuclear genes encoding MRC ...

Mitochondrial genetics: 9. mtDNA and aging
...(biological clock abnormal) phenotype characterized by slow development and increased lifespan ( Lemieux et al., 2001 ). The appare...

Mitochondrial genetics: 10. Perspectives
... is a vital component of animal metabolism, physiology, and development. Few studies have specifically addressed the C. elegans mtD...


Genomics and biology of the nematode Caenorhabditis briggsae [HTML]
Genomics and biology of the nematode Caenorhabditis briggsae
...coding RNAs 2.9. Gene families 3. Evolutionary studies 3.1. Development of the excretory duct cell 3.2. Sex determination 3.3. Development of the vulva 3.4. Development of the male tail 3.5. Thermal sensitivity 3.6. Specificatio...

...tractive model system for studying evolution of both animal development and behavior. Being a close relative of C. elegans, C. brig...

Genomics and biology of the nematode Caenorhabditis briggsae : 3. Evolutionary studies
...g of the evolution of sex-determination pathways. 3.3. Development of the vulva Nematode vulval development is an established model for comparative developmental biolo...

...f function alleles, on the other hand, do not affect vulval development. However, in C. briggsae , glp-1 is required for vulval development. Cbr-glp-1 (RNAi) animals exhibit a multivulva phenotype su...

...pecific genes in mediating some of these changes. 3.1. Development of the excretory duct cell The excretory system in C. elega...

...-1 is necessary during early embryogenesis and later in the development and proliferation of the germline whereas LIN-12 , but not ...

Genomics and biology of the nematode Caenorhabditis briggsae : 4. Genetic approach to study functions of C. briggsae genes
...nes. Some of the studies that are currently ongoing involve development of the vulva (B. Gupta, unpublished), sex determination ( H...

Genomics and biology of the nematode Caenorhabditis briggsae : 5. Tools for genetic studies
...). While the experimental protocol is in the early phase of development in C. briggsae , it has potential of rapidly isolating alle...


The cuticle [HTML]
The cuticle
...rowth by molting. It is synthesised five times, once in the embryo and subsequently at the end of each larval stage prior to m...

The cuticle : 1. Introduction
...t as the mature cuticle. Synthesis occurs five times during development, once in the embryo, then prior to molting at the end of each larval stage. Thu...

...wormbook.1.138.1 1. Introduction During post-embryonic development, Caenorhabditis elegans is enclosed within an exoskeleton c...

The cuticle : 6. Collagen biosynthetic pathway
...DPY-7 (green) and PDI-2 (red) double staining of a 1.5-fold embryo. At this stage both PDI-2 and DPY-7 are co-localised in the...

...Y-7 (green) and PDI-2 (red) double staining of an elongated embryo. At this stage DPY-7 (green) has been secreted and is prese...

...sation in early comma stage embryos. Two images of the same embryo taken at focal planes 1 μm apart. SEC-23 localises to ...


The cadherin superfamily [HTML]
The cadherin superfamily
...e processes is not known. FMI-1 is required during neuronal development consistent with the known role of the Drosophila homologue ...

The cadherin superfamily : 1. The cadherin superfamily
...both of mechanism and function within the context of animal development. The defining feature of this family is the presence of a c...

...m indicates that classic cadherins also act during neuronal development in C. elegans . Animals with reduced or absent HMR-1B funct...

...gions of contact between all cells in the pre-morphogenetic embryo (the endogenous expression of JAC-1 is not known; Nance et ...

...c cadherin function, or adhesion between cells of the early embryo does not involve cadherin function. ...

The cadherin superfamily : 2. The FAT-like cadherins: CDH-3 and CDH-4
... mutants do not display any detectable defects in epidermal development (J. Pettitt, unpublished). cdh-4 promoter reporter construc...

The cadherin superfamily : 3. FMI-1, a C. elegans FLAMINGO/STAN cadherin
...nalling. Fmi/Stan is also required for axon guidance during development of the Drosophila visual system where it functions, at leas...

...or FMI-1 in the regulation of cell polarity during neuronal development in C. elegans (G. Garriga and Y. Jin, personal communicatio...

The cadherin superfamily : 5. CDH-9/DCad96Cb
...dentified in a screen for genes expressed during pharyngeal development ( Gaudet and Mango, 2002 ). This indicates that CDH-9 may h...

The cadherin superfamily : 7. Conclusions and future prospects
...dherins will be identified through the analysis of neuronal development, as has proven to be the case for fmi-1 . Indeed lack of a ...


Genetic mosaics [HTML]
Genetic mosaics
... Robert K. Herman Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, MN 55455 USA This ch...

...tinct cells that require the gene for distinct fates during development. The gene bre-5 is used as an example of the usefulness of ...

Genetic mosaics : 2. How worm genetic mosaics are generated
...ement fails to be transmitted to a daughter cell during the development of the animal, all of the descendants of that cell, a clone...

...astomeres for genetic studies of Wnt signaling during early development ( Thorpe et al., 1997 ; Schlesinger et al., 1999 ). ...

Genetic mosaics : 3. A few examples of mosaic analysis in worms
...ression of a green marker also present on the array. During development of the embryo in A, the array was inherited by MS but not by E, its siste...

...osaic analysis has contributed important insights into worm development were recently reviewed ( Yochem et al., 2000 ; Yochem and H...

...sked by a critical requirement for the molecules earlier in development. Mosaics can in some cases uncover the foci of activity of ...

...hich encodes an ERK2-type MAP-kinase known to affect vulval development ( Lackner et al., 1994 ; Wu and Han, 1994 ). For proper vul...


Oscheius tipulae [HTML]
Oscheius tipulae
...cular biology resources 6. Developmental biology 6.1. Gonad development 6.2. Vulva development 7. Acknowledgement 8. References Abstract Oscheius tipulae ...

...resting evolutionary changes in mode of reproduction, gonad development, body size, etc. ...

Oscheius tipulae : 6. Developmental biology
...ernberg, 1996 ; Sudhaus and Hooper, 1994 ). 6.2. Vulva development Oscheius tipulae has been mostly studied for its vulva development ( see Evolution of development in nematodes related to C. elegans ). Several features make...

...mbook.1.119.1 6. Developmental biology 6.1. Gonad development Gonad development and morphology of O. tipulae are overall similar to those o...

...ancestral (Kiontke et al., in preparation). Figure 3. Vulva development in Oscheius tipulae .  Left panel. Successive stages o...

...ius tipulae .  Left panel. Successive stages of vulval development. Differences with C. elegans are highlighted in green. AC: ...


Strongyloides stercoralis: a model for translational research on parasitic nematode biology [HTML]
Strongyloides stercoralis: a model for translational research on parasitic nematode biology
...cess in parasites and its similarity to regulation of dauer development in C. elegans. Furthermore, we have developed a practical m...

Strongyloides stercoralis: a model for translational research on parasitic nematode biology : 2. Laboratory hosts for S. stercoralis
...o alternative pathways. The homogonic cycle involves direct development to the infective L3 (iL3), and heterogonic development involves development to the free-living female and, following mating, production...

...is species are capable of undergoing only one generation of development outside the host and continuous laboratory rearing requires...

...lowing repeated, but well-regulated cycles of autoinfective development whereby barren or moribund female worms are replaced by the...

...he autoinfective cycle, unique to S. stercoralis , in which development from the L1 to the autoinfective L3 (aL3) occurs within the...

Strongyloides stercoralis: a model for translational research on parasitic nematode biology : 5. Transgenesis
...hese plates are incubated at 22°C for 5-6 days to allow development to the iL3, which are collected by the Baermann funnel prot...

Strongyloides stercoralis: a model for translational research on parasitic nematode biology : 6. Neuronal ablation
...heterogonic developmental alternatives and of resumption of development by iL3, so we typically screen larvae for phenotypes follow...

Strongyloides stercoralis: a model for translational research on parasitic nematode biology : 7. Future prospects
...ect with which to build upon these advancements for further development of a forward genetic system in a parasitic nematode ( Grant...


Dauer [HTML]
Dauer
..., USA This chapter is in WormBook section: > Post-embryonic development View/Add Comments Table of Contents 1. Introduction 2. Envi...

Dauer : 1. Introduction
...mechanisms governing morphological change during organismal development as well as a parallel to obligate dauer-like developmental ...

Dauer : 5. Pheromone
...ve not emerged from the extensive genetic analysis of dauer development that has been performed to date. The availability of synthe...

Dauer : 6. Molecular characterization of signaling pathways regulating dauer arrest
...hat regulates thermosensation, chemosensation, and neuronal development ( Coburn and Bargmann, 1996 ; Coburn et al., 1998 ; Komatsu...

...equences. Furthermore, insulin-like signaling during larval development regulates dauer arrest without significantly impacting life...

...ersonal communication). INS-7 may also promote reproductive development, as ins-7 RNAi enhances dauer arrest in a daf-2(e1370) muta...

...ecules ( Oldham and Hafen, 2003 ) also promote reproductive development in C. elegans . TOR is a protein kinase activated by insuli...

Dauer : 9. Modifier screens
... potentially identify genes required for ASI specification, development, and function. Three sdf genes have been described. sdf-13 ...

Dauer : 10. The XXX cells as a site of integration of insulin-like and steroid hormone signaling
...hough the XXX cells clearly play a role in regulating dauer development, the relative weakness of the XXX laser ablation dauer arre...

Dauer : 11. Expression profiling of dauers
...te insulin-like and steroid hormone signaling during larval development in replete environments. daf-9 expression increases early a...

Dauer : 14. Future directions
... we learn about their interactions during C. elegans larval development may be germane to the interactions of similar signaling pat...


Genetic enhancers [HTML]
Genetic enhancers
...ct; , John Yochem Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, MN 55455 USA This ch...

Genetic enhancers: 1. Overview
...o become increasingly important for the elucidation of worm development and behavior. Figure 1. Synthetic interaction between mec-8...

...is not formed. Unable to feed, hatchlings arrest growth and development during the first larval stage. The mutants also have a bulb...

Genetic enhancers: 2. Dominant enhancers
... mutations in genes that act in a pathway for male tail ray development has also been reported ( Ikegami et al., 2004 ). Figure 2. ...

Genetic enhancers: 4. Enhancement of mutations in homologous genes
...edundantly-acting ligands for LIN-12 signaling during vulva development ( Chen and Greenwald, 2004 ). ...

Genetic enhancers: 5. Enhancement of mutations in functionally redundant but nonhomologous genes
...rphogenesis ( Fay et al., 2003 ; Fay et al., 2004 ), larval development ( Cui et al., 2004 ), and somatic gonad development ( Bender et al., 2004 ). Synthetic interactions with lin-35...

...Ferguson and Horvitz, 1989 ). The Muv phenotype (see Vulval development ) requires two SynMuv mutations, one in a SynMuv class A ge...

...ene. Two overlapping but nonhomologous pathways repress Muv development, which occurs only when both pathways are inactivated. Evid...

...or the mutation in gene-A has no apparent effect on growth, development, and fertility of worms grown on a bacterial lawn (left pla...

Genetic enhancers: 6. Recessive enhancers of non-null mutations
... genes previously known to play essential roles in germline development were identified as well as mutations in five new genes ( Qi...

...RNA-directed RNA polymerase that functions both in germline development and in the process of RNA interference ( Smardon et al., 20...


Chemosensation in C. elegans [HTML]
Chemosensation in C. elegans : 1. Chemosensation and its regulation by ciliated sensory neurons
...4 ; Qin et al., 2001 ). A transcriptional program for cilia development is controlled by daf-19 . which encodes an X box transcript...

... temperatures, high density, and limiting food modify their development to form long-lived and stress-resistant dauer larvae ( Gold...

...he same cues to decide whether to recover and resume normal development. Both density and food are sensed by ciliated sensory neuro...

...ocrine signal to prevent dauer formation and promote normal development ( Ren et al., 1996 ; Schackwitz et al., 1996 ). Expression ...

Chemosensation in C. elegans : 2. Signal transduction in chemosensation
...pressed in AWA, AWC, and non-neuronal cells, affects vulval development and olfactory plasticity by modulating MAPK signaling ( Bat...

...xcessive sensory axon outgrowth occurs during postembryonic development, while the initial outgrowth of the sensory axons occurs in...

...le the initial outgrowth of the sensory axons occurs in the embryo ( Coburn et al., 1998 ). Since TAX-2 and TAX-4 are primaril...

...tein signaling. daf-11 , which is required to prevent dauer development, is expressed in the ASI and ASJ neurons that regulate daue...


The measurement and analysis of age-related changes in Caenorhabditis elegans [HTML]
The measurement and analysis of age-related changes in Caenorhabditis elegans
...0 USA This chapter is in WormBook section: > Post-embryonic development View/Add Comments Table of Contents 1. Measurements of age-...

...anges. 1.2. Distinguishing age-related changes pertinent to development and senescence. 1.3. Approaches to measuring age-related ch...

The measurement and analysis of age-related changes in Caenorhabditis elegans : 1. Measurements of age-related changes
.... 1.2. Distinguishing age-related changes pertinent to development and senescence. Given that age-related changes are widespre...

...aw a distinction between age-related changes that represent development and growth and age-related changes that represent senescenc...

...int in the life cycle when the change occurs. Events in the embryo will typically be developmental, whereas changes in older a...

...However, there is no time in midlife that clearly separates development from aging, and it is possible that developmental changes a...


Carbohydrates and glycosylation [HTML]
Carbohydrates and glycosylation : 1. Introduction
...ns have made to our understanding of the role of glycans in development. ...

Carbohydrates and glycosylation : 2. Glycosylation pathways in C. elegans
...ns. Our understanding of the role of glycoconjugates in the development of C. elegans to date is largely due to the identification ...

Carbohydrates and glycosylation : 3. Glycosaminoglycans
...the expansion of the extracellular spaces in the C. elegans embryo and vulva; among other possibilities ( Hwang et al., 2003 )...

...legans : rib-1 and rib-2 . rib-2 is essential for embryonic development ( Morio et al., 2003 ) and encodes a unique α 1,4-N-ac...

Carbohydrates and glycosylation : 5. O-linked glycans
...mals, the OGT gene is essential for embryonic and stem cell development and produces multiple transcripts ( Hanover et al., 2003 ; ...

Carbohydrates and glycosylation : 6. Glycolipids
...rops expressing Cry proteins to control insect pests is the development of resistance among target populations. Mutations in the br...

Carbohydrates and glycosylation : 7. Chitin
...chs-1 and chs-2 play different and non-overlapping roles in development. ...

Carbohydrates and glycosylation : 8. Conclusions
...t how the expression of these molecules is regulated during development and differentiation. C. elegans and small model organisms a...

...tions of glycan structure can be analyzed in the context of development and differentiation. Sensitive techniques for glycan analys...

...ng model system for investigating basic roles of glycans in development. ...


Complementation [HTML]
Complementation: 1. A simple test for assigning a mutation to a genetic locus
...3 is required in at least four processes, viability, vulval development, male spicule development and hermaphrodite fertility. lin-3 alleles exhibit a differ...

...t three different processes; vulval induction, male spicule development and hermaphrodite fertility ( Ferguson and Horvitz, 1985 ; ...

... of another allele, n1058 , exhibit defects in male spicule development, hermaphrodite fertility, viability, and only very weakly, ...

Complementation: 2. When the complementation test is not so simple
...disrupts vulval induction, having no effect on male spicule development, while the n1058 mutation only weakly disrupts vulval induc...

...srupts vulval induction, but strongly disrupts male spicule development ( Liu et al., 1999 ). These homozygous phenotypes suggest t...

...ic enhancer, which is required for LIN-3 function in vulval development, but would not affect LIN-3 function in other tissues ( Hwa...

...a temporally and spatially regulated manner during germline development, and these requirements are reflected in the range of mutan...


X-Chromosome dosage compensation [HTML]
X-Chromosome dosage compensation: 3. Dosage compensation proteins assemble onto hermaphrodite X chromosomes to regulate
...both X chromosomes of hermaphrodites. Shown is a multi-cell embryo stained with the DNA intercalating dye DAPI (blue) and an a...

... the holocentric mitotic chromosomes. Shown is a two-celled embryo stained with the DNA dye DAPI (red) and antibodies to SMC-4...

X-Chromosome dosage compensation: 4. Sex-specific targeting of the dosage compensation complex to hermaphrodite X chromosomes
... et al., 1988 ). When xol-1 is active (in XO embryos), male development ensues; when xol-1 is inactive (in XX embryos) hermaphrodit...

...nsues; when xol-1 is inactive (in XX embryos) hermaphrodite development ensues, including the activation of dosage compensation ( R...

...s. SDC-2 also activates the hermaphrodite program of sexual development by repressing the male-specific sex-determination gene her-...

X-Chromosome dosage compensation: 6. Recruitment of the dosage compensation complex for gene-specific versus
...ific factor to initiate the hermaphrodite program of sexual development came in part from the observation that overexpression of SD...

...lop as hermaphrodites. SDC-2 activates hermaphrodite sexual development by binding directly to regulatory regions of the male sex-d...


Homologs of the Hh signalling network in C. elegans [HTML]
Homologs of the Hh signalling network in C. elegans
...n cell proliferation and survival, and in patterning of the embryo, limb buds and organs. In C. elegans, this pathway has unde...

Homologs of the Hh signalling network in C. elegans : 1. Introduction
...e collectively proven to be key effectors in patterning the embryo, limb buds and neural tube, organ formation, cell prolifera...

Homologs of the Hh signalling network in C. elegans : 2. Hh pathway components in C. elegans
... role does the Hh/Ptc signalling pathway play in C. elegans development? To address this question, it is first reasonable to ask wh...

...evel to control sexual cell fate decisions and male gonadal development ( Hodgkin, 1983 ; Mathies et al., 2004 ), although there is...

Homologs of the Hh signalling network in C. elegans : 4. Modification and function of Hh-r genes
...ary results indicate that they also play important roles in development. For example, wrt-5 functions in the hypodermis during embr...

Homologs of the Hh signalling network in C. elegans : 6. Role of PTC proteins in C. elegans
...t carry multinucleate oocytes and sperm. In other respects, development of a ptc-1 mutant is essentially wild type. In situ hybridi...

...nerated membrane furrows leads to a loss of cell autonomous development in the germline syncytium, as evidenced by the presence of ...


Reverse genetics [HTML]
Reverse genetics : 1. Introduction to reverse genetics in C. elegans
...hniques, a gene's function is altered and the effect on the development or behaviour of the organism is analysed. Reverse genetics ...

Reverse genetics : 4. RNAi by soaking
...orms) are examined for phenotypes. In many cases, the first embryo laid is already fully affected by RNAi. L1-soaking : To per...

...ly affected by RNAi. L1-soaking : To perform post-embryonic development specific knockdowns, L1 larvae are soaked in a dsRNA soluti...

... During soaking (which is done in the absence of food), the development of the L1 larvae is arrested but the RNAi response continue...

... to suppress gene function from the onset of post-embryonic development. 4.2.1. L4-soaking Day 1: Soaking In a 200μl PCR ...

Reverse genetics : 5. RNAi feeding on agar plates
...orms. Also, as many genes are required at multiple times in development, different phenotypes may be seen when using L1s compared t...

Reverse genetics : 6. RNAi feeding in liquid culture (96 well format)
...owledgements We thank members of the lab for input into the development of this protocol and comments on the manuscript. We also gr...

Reverse genetics : 7. Construction and screening of deletion mutant libraries to generate C. elegans gene knockouts
...t of the F2 animals do not run out of food and arrest their development until after they are past the L1 stage. Because these older...


DNA repair [HTML]
DNA repair
...lay essential roles in DNA replication, cell cycle control, development, meiosis and mitosis. To date, no obvious DNA damage-induce...

DNA repair : 1. Overview
...F43G6.1 Rad – Chromosome structure coh-2 F10G7.4 Late embryo or larval arrest or HIM – scc-3 F18E2.3 Embryonic let...

DNA repair : 2. Radiation sensitivity of C. elegans
...y embryogenesis. The UV hypersensitivity of the early stage embryo might be due in part to the rapid cell proliferation and th...

DNA repair : 3. DNA damage checkpoints in the germ line
... checkpoints in the germ line In contrast to the developing embryo, activation of the checkpoint in the germ line results in o...

...ne, but not for programmed cell death occurring during worm development nor physiological (radiation-independent) germ cell death (...


Synaptogenesis [HTML]
Synaptogenesis
...enesis * Yishi Jin § Department of Molecular, Cell and Development Biology, Howard Hughes Medical Institute, University of Cal...

...pter is in WormBook section: > Neurobiology and behavior >> Development View/Add Comments Table of Contents 1. Introduction 2. Visu...

...cognition 6. Synaptic ECM 7. Activity-dependence of synapse development 8. Perspectives 9. Acknowledgement 10. References Abstract ...

Synaptogenesis : 4. Presynaptic assembly
...ily (see Asymmetric cell division and axis formation in the embryo ). SAD-1 localization intermingles with SV cluster, but doe...

...ed p38 MAP kinase mediated signaling cascade in presynaptic development, and further suggest that activity of this MAP kinase casca...

Synaptogenesis : 7. Activity-dependence of synapse development
...1895/wormbook.1.44.1 7. Activity-dependence of synapse development In vertebrate nervous system, synaptic activity plays a cru...

... muscle arms ( Loria et al., 2004 ). In unc-122 mutants NMJ development is morphological normal, the GABAergic DVB neuron displays ...


Specification of the nervous system [HTML]
Specification of the nervous system
...pter is in WormBook section: > Neurobiology and behavior >> Development View/Add Comments Publication History version 1 latest vers...

Specification of the nervous system : 2. Neuronal vs. non-neuronal
...fined precursor cells in specific regions of the developing embryo, a brief look at the lineage of C. elegans illustrates, str...

...es ( Figure 1 ). However, in analogy to vertebrate neuronal development ( Hemmati-Brivanlou and Melton, 1997 ), it is conceivable t...

Specification of the nervous system : 4. Neuron class specification
...af-19 transcription factor was found to be required for the development of specific differentiated aspects of all ciliated sensory ...

...gene regulatory factors that act at late stages of neuronal development, most often during terminal differentiation of specific sub...

Specification of the nervous system : 7. Concepts in neuronal fate specification in C. elegans
...evels; for example, unc-86 works at different stages in the development of neuroblast lineage and presumably interacts with distinc...

... (circles indicate the expression of the character). During development and/or evolution (indicated by arrows), sequential repressi...


Introduction to the germ line [HTML]
Introduction to the germ line: 1. An unbroken chain: the germ line
...n chain, from generation to generation. C. elegans germline development can be conceptually divided into three phases: specificatio...

...al stages, the germ line proliferates and undergoes meiotic development. During the remainder of reproductive life the mature germ ...

...fe the mature germ line continues proliferation and meiotic development, and produces gametes. Chapters in the Germ line section fo...

...ters in the context of the temporal progression of germline development in hermaphrodites ( Figure 1 ). Figure 1. Cartoon represent...

Introduction to the germ line: 2. Embryogenesis and early L1: germline specification
... not sufficient for specifying the germ cell fate, germline development is severely impaired in the absence of their constituent pr...


Biology and genome of Trichinella spiralis [HTML]
Biology and genome of Trichinella spiralis : 1. Introduction
...e muscle fibers, they encyst ( Figure 2 , panel B), undergo development, become infective within 15 days and remain for months to y...

... under debate ( Kozek, 2005 ). Furthermore, the embryologic development of the Dorylaimia species involves anterior positioning of ...

...bed for Trichinella spp., similarities between the arrested development displayed by muscle larvae and C. elegans dauer larvae have...

Biology and genome of Trichinella spiralis : 2. Molecular and cellular interactions with the host
...lso unresolved. A significant advance in this regard is the development of an in vitro culture system for the intestinal phase of T...

... al., 2002 ; ManWarren et al., 1997 ). This system supports development to the adult stage and offers a valuable tool to answer the...

...se cells. Inhibition of both invasion and in vitro parasite development was accomplished with antibodies against a glycan molecule ...

...cle cells may be adaptive for nutrient acquisition, growth, development and/or altered detection by the host immune system. Worthy ...

Biology and genome of Trichinella spiralis : 5. Current genome sequencing project for Trichinella spiralis
... marked by important changes in animal anatomy, physiology, development or behavior. In addition, the T. spiralis genome sequence w...


Genetic suppression [HTML]
Genetic suppression
...hierarchies. In C. elegans biology, the processes of muscle development, vulva formation and sex determination have provided remark...

Genetic suppression : 8. Nonsense suppression
... is sufficient to provide enough activity for normal sexual development, so weak suppressors can be recovered. As indicated in Tabl...

... gene expression, amber suppressors have been useful in the development of methods for microinjection and transformation of C. eleg...

Genetic suppression : 9. Extragenic suppression by modified splicing
...ts when homozygous: sup-6 homozygotes arrest in late-larval development, and 50 % of sup-39 homozygotes die as embryos. In addition...

Genetic suppression : 13. Bypass suppression
...llustrate this effect ( Zhang and Emmons, 2000 ). In normal development, pal-1 expression in certain tail cells depends on a positi...

Genetic suppression : 16. Suppression by modulators of activity
...h is involved in numerous important signaling events during development (see LIN-12/Notch signaling in C. elegans ). Suppression ex...

Genetic suppression : 17. Suppression by epistasis
...egulatory pathways in C. elegans . Many important events in development depend on signal transduction events, whereby an inductive ...

..., the feminizing transcription factor TRA-1 promotes female development, and the membrane receptor TRA-2 prevents inhibition of TRA...


Embryoinsitu.pdf [preprint PDF]
Embryoinsitu.pdf
...r 75 sec. 3. Wash the embryos 3 times with EH buffer (Embryo Handling buffer). 4. Wash the embryos with Basal EH buf...

...ne coated 8-well slides. 7. Dispense 5 ul/well of the embryo suspension into the buffer at each well. 8. Let stand f...

...ee reagetnts) twice for 5 min each at r.t.. 6. Colour development i. Mix 180 ul of NBT and 140 ul of BCIP in 40 ml of st...

...7.2 and autoClave PBT PBS + 0.1%Tween 20 EH buffer (Embryo Handling buffer) mannitol 0.3M Hepes pH 7.2 50mM NaCl...


Genetic balancers [HTML]
Genetic balancers : 2. Types of balancers
...a large percentage of aneuploid progeny (10/16) that arrest development ( Figure 2 ) as embryos or early larvae ( Adames, K.A., Gaw...

... representing aneuploid progeny, all of which arrest during development, are shaded. All wild-type progeny are heterozygous for the...

...loss of certain free duplications from somatic cells during development has made possible mosaic analysis of the gene function ( He...

Genetic balancers : 3. Practical considerations of balancer use
...n. All other cross-progeny are aneuploid, and arrest during development. 3.4. Accumulation of spontaneous mutations Rosenbluth...

... phenotypically undetectable in a lethal that arrests as an embryo. These unrecognized secondary mutations may confound analys...

Genetic balancers : 5. A field guide to balancers
...istics: Egg hatching nearly 100%. About 20% of these arrest development as early larvae, and are presumably duplication homozygotes...


Genomic overview of protein kinases [HTML]
Genomic overview of protein kinases : 3. Kinase evolution
...ypical H11   0 1 Apoptosis?   CAMK CAMKL HUNK 0 1 Development Mammary gland development CAMK Trio   0 6 Muscle Human adds kinase to conserved ...

... morphogenesis; immunity   TK Ret   1 1 Immunity, development Growth factor receptor TK Sev   1 1 Morphogenesis &nbs...

... LISK LIMK 1 2 Cytoskeletal   TKL LISK TESK 1 2 Testis development   Human Atypical Alpha ChaK 0 2 Neuronal Human adds ki...

...  0 2 Cytoskeletal   Other NKF5   0 2 Testis development?   TK Axl   0 3 Cell growth; adhesion   TK L...

Genomic overview of protein kinases : A. Appendix A: Classification of worm kinases
... F23C8.8 Asymmetric cell division and axis formation in the embryo Microtubule affinity regulating kinase CAMK CAMKL MELK 1 Ne...


Introduction to sex determination [HTML]
Introduction to sex determination
... K. Herman § Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, MN 55455 USA This ch...

...self-fertilization are encased in an egg shell and initiate development within the uterus of the hermaphrodite. When they reach abo...

...st differences between the sexes arise during postembryonic development through different patterns of cell lineage. Surprisingly, t...

...er of this section, Scott Emmons describes those aspects of development that are specific to the male. Much of the interesting male...


Introduction to signal transduction [HTML]
Introduction to signal transduction
...sduction in all animals, and defined their diverse roles in development and cell physiology. More recently, the development of "reverse" genetic approaches that exploit the genomic se...

...ny different roles for RTK-Ras-MAPK signaling in C. elegans development. She has also described how genetic studies of Ras-mediated...

... roles of TGF- β ; signaling in regulating dauer larva development, body size, and axon guidance (see TGF- β signaling )....

... a comprehensive overview of the many aspects of C. elegans development that are controlled by Wnt signaling (see Wnt signaling ). ...


The biology and genome of Heterorhabditis bacteriophora [HTML]
The biology and genome of Heterorhabditis bacteriophora : 5. Symbiosis
...red for P. temperata to support H. bacteriophora growth and development (discussed below) and is unable to compete with wild-type b...

.... luminescens or P. temperata as a substrate for growth and development ( Akhurst et al., 1996 ) and is very sensitive to contamina...

...ition, the primary form is required for nematode growth and development. The primary form is isolated from IJs and infected insects...

...c exchange and neither mutant supported nematode growth and development ( Bintrim and Ensign, 1998 ). The mutant cip strains were p...

The biology and genome of Heterorhabditis bacteriophora : 7. Genetics and molecular biology of H. bacteriophora
...nome project. The genome and a physical map, along with the development of RNAi and transformation, should facilitate the development and use of genetic and molecular tools in H. bacteriophora....

...( Nugent et al., 1996 ). Despite these encouraging results, development of genetics and molecular biology in Heterorhabditis is sti...


Germline survival and apoptosis [HTML]
Germline survival and apoptosis : 2. Methods to assess germline apoptosis
...requency, as can be done for somatic apoptotic death during development ( Schwartz, 2007 ). Figure 1. The C. elegans hermaphrodite ...

Germline survival and apoptosis : 4. Functions, regulation, and conservation of physiological germ cell apoptosis
...tosis lip-1 6 MAPK ( MPK-1 ) phosphatase Accelerated oocyte development and partial embryonic lethality; reduced germline prolifera...

Germline survival and apoptosis : 5. DNA damage-induced germ cell apoptosis
...ion stress, and also for an essential function in germ cell development that is most likely linked to DNA replication ( Garcia-Muse...

...-2(qm37) was found in a screen for mutations that slow worm development and lead to an extended life span ( Benard et al., 2001 ; H...

Germline survival and apoptosis : 6. cep-1 (p53/p63) and the regulation of DNA damage-induced germ cell apoptosis
...ional repressor, which controls several aspects of germline development, negatively regulates cep-1 expression ( Schumacher et al.,...

Germline survival and apoptosis : 10. Conclusions
...osis is induced by an important control point during oocyte development, and a novel and as yet unknown mechanism for activating th...

Germline survival and apoptosis : 11. Acknowledgements
...inhard Family Foundation (PRB), a Cancer Research UK career development award fellowship (AG) and an AICR project grant (AG). We th...


Intracellular trafficking [HTML]
Intracellular trafficking
...odified in the context of a multicellular animal undergoing development to produce diverse cell types such as muscles, nerves, and ...

Intracellular trafficking : 1. The endocytic pathway
...tly labeled BSA or dextrans. These observations allowed the development of a pulse-chase type assay of coelomocyte endocytosis ( Zh...

Intracellular trafficking : 3. General secretion
... thought to form by secretion of eggshell components by the embryo itself shortly after fertilization. Several other proteins ...

...from heterozygous mothers. Normal progression through early development presumably relies upon maternally derived SEC-23 . RNAi exp...

...experiments revealed a requirement for sec-23 during larval development, particularly during molting. Adults depleted of SEC-23 by ...


Behavior [HTML]
Behavior : 1. Introduction
...al introduction. New behavioral assays are constantly under development and old assays are often revised. Your suggestions, comment...

...g status and history. Animals that have been starved during development or animals that have not gone through the dauer stage can h...

...h many assays can be used on larval animals. Nervous system development is largely complete by the end of the L1 stage, but some de...

Behavior : 6. Thermal responses
...o-tail distance of a swimming worm using the LabVIEW Vision Development Module. Since the software works best with a high-contrast ...

Behavior : 9. Egg-laying, males and mating
...s thus staged should be within a few hours of each other in development. Put the staged animals back in the incubator, and count af...

...meaningful if you show that your strain undergoes embryonic development at about the same rate as does N2. Pick a few early 1 or 2 ...


Nematode genome evolution [HTML]
Nematode genome evolution : 3. The evolution of chromosome number and structure
...en C. elegans ' ancestor switched from a regulative mode of development to a deterministic lineage-driven mode. They suggest that p...

...III. Trichinella spiralis probably has a regulative mode of development, but it is not yet known whether its HOX genes are clustere...

...ered. However, even though Brugia malayi has lineage-driven development, most of its HOX cluster seems to have been preserved intac...

...e "X-signal elements" whose dosage determines the sex of an embryo ( Meyer, 2000 ). X-autosomal translocations may also be del...


Genetic mapping and manipulation: Chapter 1-Introduction and basics [HTML]
Genetic mapping and manipulation: Chapter 1-Introduction and basics : 3. Types of mutations
... defects in the pattern of cell divisions that occur during development. Consult WormBase or other sources for specifics concerning...

Genetic mapping and manipulation: Chapter 1-Introduction and basics : 5. Feeding, growing, and maintaining worms
...t 20 ° C for a fertile adult to develop from a one-cell embryo. At 15 ° C this process takes about twice as long, and ...

...ty much the only way to control the rate of worm growth and development. Higher temperatures (20 ° C to 25 ° C) can further...

...(20 ° C to 25 ° C) can further expedite the rate of development but can cause a drop in fertility and poor health, especial...


The phylogenetic relationships of Caenorhabditis and other rhabditids [HTML]
The phylogenetic relationships of Caenorhabditis and other rhabditids: 1. Introduction
...represented by strain CB5161 among others; see Evolution of development in nematodes related to C. elegans ). The selection of appr...

The phylogenetic relationships of Caenorhabditis and other rhabditids: 2. Rhabditid phylogeny
...ed as a satellite model organism for comparative studies of development (see Evolution of development in nematodes related to C. elegans ; Sommer, 2000 ) and mol...

...lite model system, Pristionchus pacificus (see Evolution of development in nematodes related to C. elegans ). Specifically, there a...

The phylogenetic relationships of Caenorhabditis and other rhabditids: 4. Resources for comparative biology using rhabditids
...with respect to mechanisms and patterns of gene expression, development and behavior - requires analysis in the context of the livi...


Control of oocyte meiotic maturation and fertilization [HTML]
Control of oocyte meiotic maturation and fertilization
...2. Control of oocyte meiotic maturation 2.1. Oocyte growth, development, and differentiation 2.2. Oocyte meiotic maturation 3. Cont...

Control of oocyte meiotic maturation and fertilization : 2. Control of oocyte meiotic maturation
...ntrol of oocyte meiotic maturation 2.1. Oocyte growth, development, and differentiation 2.1.1. Pachytene progression and ...

...nt the translation of proteins needed for oocyte growth and development in early meiotic prophase. The isolation of GLD-1 mRNA targ...

...precipitation identified candidate genes involved in oocyte development or function ( Lee and Schedl, 2001 ). 2.1.2. Oocyte gr...

Control of oocyte meiotic maturation and fertilization : 3. Control of ovulation
... 2004 ) as well as genes involved in sheath or spermathecal development ( Kostic et al., 2003 ; Aono et al., 2004 ). In the future,...


Pristionchus pacificus [HTML]
Pristionchus pacificus : 2. Developmental biology
...etail in P. pacificus was vulva formation (see Evolution of development in nematodes related to C. elegans ). When compared to C. e...

...elated to C. elegans ). When compared to C. elegans , vulva development in P. pacificus involves a set of evolutionary modification...

...-cell interactions during vulva formation (see Evolution of development in nematodes related to C. elegans ). P. pacificus vulva de...

...cess ( Zheng et al., 2005 ). Besides vulva formation, gonad development ( Rudel et al., 2005 ), sex determination ( Pires-daSilva &...


Anthelmintic drugs [HTML]
Anthelmintic drugs : 3. Approaches for the study of anthelmintics in C. elegans
...ificant, ideally quantifiable, change in the worm's growth, development, metabolism, and/or behaviour. Pharmacokinetic consideratio...

Anthelmintic drugs : 4. Classes of anthelmintic drugs
...and its discovery enthused other companies to invest in the development of ivermectin analogues which include moxidectin, milbemyci...

...ntrations ( Bull et al., 2007 ). The effects include slowed development, inhibition of pharyngeal pumping, decreased locomotion (fo...

...ibitory effects of emodepside on C. elegans is an important development in this field which in the light of ivermectin resistance h...

... The effect of nitazoxanide has been examined on growth and development of C. elegans ( Fonseca-Salamanca et al., 2003 ). After sev...


Biochemistry and molecular biology [HTML]
Biochemistry and molecular biology : 2. Tandem immuno-affinity purification
...ociated protein complexes to be purified from whole worm or embryo extracts by tandem immuno-affinity ( Figure 2A ; Polanowska...

...m. Lysates can also be prepared from embryos: Resuspend the embryo pellet in twice the pellet volume of homogenization buffer ...


Immunohistochemistry [HTML]
Immunohistochemistry : 1. Introduction
... of a protein within the cells of the gonad, the very early embryo, or the intestine, then the appropriate tissue can be disse...

...ted donkey anti-mouse. (B) Talin was labeled in a wild-type embryo with rabbit B547 anti-talin (gift of Bob Barstead and Gary ...


Profiling C. elegans gene expression with DNA microarrays [HTML]
Profiling C. elegans gene expression with DNA microarrays
...d to investigate changes in gene expression associated with development ( Hill et al., 2000 ; Jiang et al., 2001 ), aging ( Golden ...

Profiling C. elegans gene expression with DNA microarrays : 1. Planning your experiment
...the organism. 1.2.2. Sorting cultured cells The recent development of embryonic primary cell culture techniques for C. elegans...

...nity to profile gene expression in specific cell types. The development of cells expressing specific differentiation markers for ne...

...gene expression in a particular tissue type that occur with development, aging, or changes in environmental conditions. Clearly, th...


Introduction to nematode evolution and ecology [HTML]
Introduction to nematode evolution and ecology : 1. A model system evolves
...or building a fundamental understanding of animal genetics, development and behavior, but it is also rapidly becoming a respectable...

...important for comparative analyses of behavior, morphology, development, molecular mechanisms, and genomics. A phylogeny allows the...

...nome organization and chromosome architecture. Evolution of development in nematodes related to C. elegans provides an overview of ...

...cts of neurobiology and behavior, innate immunity, germline development and sex determination, senescence and developmental timing....


Sperm motility and MSP [HTML]
Sperm motility and MSP : 1. Spermatogenesis
...8.1 1. Spermatogenesis A detailed description of sperm development, or spermatogenesis, can be found in the WormBook chapter b...

..., which contains components that are not needed for further development or sperm function. An extracellular signal induces the proc...

...nesis and MSP.  Shown are various stages in sperm cell development. In primary spermatocytes, MSP (red) begins to assemble as ...

Sperm motility and MSP : 6. Reconstituted MSP polymerization system
...nce in our understanding of MSP-based motility has been the development of a reconstituted cell-free filament assembly system ( Ita...


The sensory cilia of Caenorhabditis elegans [HTML]
The sensory cilia of Caenorhabditis elegans
...ersity; Burnaby, BC Canada V5A 1S6 2 Center of Genetics and Development; University of California at Davis, Davis, CA 95616 USA Thi...

The sensory cilia of Caenorhabditis elegans: 5. Transcriptional regulation of cilium morphogenesis
...nesis in other organisms. In mice, RFX3 is required for the development of embryonic nodal cilia ( Bonnafe et al., 2004 ). Similarl...

...rly, an RFX protein in Drosophila has been connected to the development of ciliated structures in sensory neurons ( Dubruille et al...

The sensory cilia of Caenorhabditis elegans: 7. Understanding C. elegans ciliary functions through ciliary mutant analysis
...saw the value of this organism for studying the genetics of development and nervous system function. To this end, he and many of hi...


Obesity and the regulation of fat metabolism [HTML]
Obesity and the regulation of fat metabolism
... nhr-49 / mdt-15 4.3. TOR, AMPK, and hexosamine pathways 5. Development of fat storage capacity 6. Lipid uptake/transport 7. Neuroe...

Obesity and the regulation of fat metabolism : 3. Metabolic pathways
...ation stage (see aging and dauer chapters in Post-embryonic development section of WormBook). Increased fat accumulation and altere...

Obesity and the regulation of fat metabolism : 5. Development of fat storage capacity
...10.1895/wormbook.1.130.1 5. Development of fat storage capacity During mammalian adipogenesis, horm...

Obesity and the regulation of fat metabolism : 8. Perspectives
...ng from metabolism, transcription and signaling to neuronal development and behavior. Deciphering neuronal circuits that coordinate...


Acetylcholine [HTML]
Acetylcholine : 2. Cholinergic pharmacology and drug-resistant mutants
...ade et al., 2005 ). An alternative is to measure growth and development in the presence of aldicarb, i.e., “chronic” ex...

Acetylcholine : 4. Choline transport
...imals homozygous for tm373 are viable, and their growth and development appear to be normal. However, although cho-1 mutants have l...

Acetylcholine : 5. Acetylcholinesterases
...e-2 ; ace-3 ; ace-1 triple mutant is lethal - its embryonic development is apparently normal, but many of the animals do not hatch,...

Acetylcholine : 10. Additional aspects of cholinergic biology
...e nicotinic receptor agonist DMPP slowed overall C. elegans development during the L2 stage but did not affect the molt timing, so ...


Male mating behavior [HTML]
Male mating behavior : 3. Response behavior
... and 2 sensory neurons RnA and RnB (n = the ray number; for development of rays, see Male development ). The dendritic processes of RnA and RnB extend down the l...

Male mating behavior : 5. Vulva location
...Insulin receptor Lipton et al., 2004   glp-1 Germ cell development     spe-26 Germ cell development   Response osm-5 Cilium structure genes Barr and Stern...


A quick tour of nematode diversity and the backbone of nematode phylogeny [HTML]
A quick tour of nematode diversity and the backbone of nematode phylogeny : 1. Nematodes are highly diverse, complex and specialized metazoans
...is. A very recent case in point is the discovery that early development in Tobrilus diversipapillatus passes through a classical me...

A quick tour of nematode diversity and the backbone of nematode phylogeny : 4. Major features of Enoplea
...al within nematodes, such as a highly indeterminate mode of development ( Justine, 2002 ) and retention of the nuclear envelope in ...

A quick tour of nematode diversity and the backbone of nematode phylogeny : 5. Features and diversity of Chromadorea
..., F ). A key adaptation within the order Rhabditida was the development of a chemically impermeable cuticle, which clearly contribu...

...). Within the order Rhabditida, a major factor has been the development of a modified juvenile stage specifically adapted to long-t...


Alternative splicing in C. elegans [HTML]
Alternative splicing in C. elegans: 1. The Importance of alternative splicing
...plicing occurs in C. elegans and plays an important role in development. This chapter builds upon a previous review in C. elegans I...

Alternative splicing in C. elegans: 4. Constitutive splicing factors in C. elegans
... Family members are expressed in all cells at all stages of development ( Kawano et al., 2000 ). There is functional overlap among ...

...te and activity level of these proteins change during early development ( Sanford and Bruzik, 1999a ; Sanford and Bruzik, 1999b ). ...

Alternative splicing in C. elegans: 7. Regulation of alternative splicing in C. elegans
...-1 , is expressed only in muscle cells and is essential for development ( Milne and Hodgkin, 1999 ). ...


The sensory cilia of Caenorhabditis elegans [HTML]
The sensory cilia of Caenorhabditis elegans
...ersity, Burnaby, BC Canada V5A 1S6 2 Center of Genetics and Development, University of California at Davis, Davis, CA 95616 USA Thi...

The sensory cilia of Caenorhabditis elegans: 5. Transcriptional regulation of cilium morphogenesis
...nesis in other organisms. In mice, RFX3 is required for the development of embryonic nodal cilia ( Bonnafe et al., 2004 ). Similarl...

...rly, an RFX protein in Drosophila has been connected to the development of ciliated structures in sensory neurons ( Dubruille et al...

The sensory cilia of Caenorhabditis elegans: 7. Understanding C. elegans ciliary functions through ciliary mutant analysis
...saw the value of this organism for studying the genetics of development and nervous system function. To this end, he and many of hi...


Neuropeptides [HTML]
Neuropeptides : 4. Identification of putative neuropeptide genes
...head neurons, VNC, 1 ant. pharyngeal neuron, 1 tail neuron, embryo, intestine   1, 2, 3, 4 nlp-22 T24D8.3 X SIAIGRAGFRPG ...

Neuropeptides : 7. Neuropeptide function
... INS-19 may signal through DAF-2 to affect other aspects of development. Overexpression of ins-4 or ins-6 can suppress or partially...


Reporter gene fusions [HTML]
Reporter gene fusions : 1. General considerations
...periments, mosaic analysis and antibody staining. For early embryo studies, in situ hybridization can also be informative. 1.1...

Reporter gene fusions : 2. Protocols
...ances in the molecular understanding of GFP have led to the development of reconstituted GFP expression systems. Partial GFP peptid...


GABA [HTML]
GABA : 3. GABA neurons in nematodes
... data had indicated that GABA could be involved in neuronal development, specifically in cell proliferation, cell migration, neurit...

...in mice have demonstrated that GABA does not play a role in development ( Ji et al., 1999 ). Careful analysis of the unc-25 mutant ...

...nc-25 mutant phenotype failed to reveal defects in neuronal development. Axon morphology of GABA neurons is normal in adult unc-25 ...


Genetic mapping and manipulation: Chapter 8-Dominant mutations [HTML]
Genetic mapping and manipulation: Chapter 8-Dominant mutations : 4. Different types of dominant mutations
...-1 and imagine it is an enzyme whose activity promotes head development. Assume that normal levels of dom-1 activity result in norm...

... that normal levels of dom-1 activity result in normal head development and any dom-1 activity above normal levels results in a spi...

...en dom-1 activity gets to the threshold required for normal development. A point mutation that makes this serine phosphorylation im...


Gene duplications and genetic redundancy in C. elegans [HTML]
Gene duplications and genetic redundancy in C. elegans: 1. Gene duplications and genetic redundancy
... greater number of duplicates than those expressed early in development. This was found to be true in both the C. elegans and C. br...

...nt one is the fbf-1 / fbf-2 gene pair involved in germ line development in C. elegans . fbf-1 and fbf-2 encode closely related RNA ...

...xpressed in intestinal cells and is required for intestinal development ( Fukushige et al., 1998 ). While it is clear that elt-4 is...


Caenorhabditis briggsae methods [HTML]
Caenorhabditis briggsae methods : 1. The Caenorhabditis briggsae model system
...03 ). From this point, these two model systems diverge. The development, behavior, and physiology of C. elegans have been character...

Caenorhabditis briggsae methods : 2. Natural history and population biology
...oenvironments. In necromenic associations, nematodes resume development and reproduce in the bacterial bloom that occurs following ...

Caenorhabditis briggsae methods : 5. Online resources for C. briggsae genetics and genomics
...line resources for C. briggsae genetics and genomics As the development of C. briggsae as a model system is in its infancy, informa...


Pristionchus pacificus genetic protocols [HTML]
Pristionchus pacificus genetic protocols : 1. Introduction
.... pacificus has distinctive body morphology and diverges in development both at the cellular and molecular level when compared to C...

...lecular level when compared to C. elegans (see Evolution of development in nematodes related to C. elegans ). Despite these differe...

Pristionchus pacificus genetic protocols : 2. Protocols
...with the transformer phenotype. For genes involved in vulva development, however, the efficiency of the RNAi seems to be much lower...


Neurophysiological methods in C. elegans: an introduction [HTML]
Neurophysiological methods in C. elegans: an introduction
... channels on neuronal function has been a relatively recent development. This chapter presents a compendium of protocols and techni...

...y genes with critical functions in such processes as neural development, sensory perception, and motor activity. Because of the eas...

Neurophysiological methods in C. elegans: an introduction : 3. In vitro physiology
...used in vitro and in vivo ( Samuel et al., 2003 ). With the development of novel neural indicator molecules as well as new techniqu...


Basement membranes [HTML]
Basement membranes
... provided insights into their assembly and functions during development. Immuno- or GFP-tagged localization studies have shown that...

...how these structures are assembled and how they function in development. ...

Basement membranes : 7. SPARC/Osteonectin
...types suggests that OST-1 has multiple functions throughout development. ...


Ecology of Caenorhabditis species [HTML]
Ecology of Caenorhabditis species: 2. Associations with other animals
... animal and wait for their carrier to die. They then resume development and propagate on the decomposing cadaver. Typically, dauer ...

...e they disembark from the living or dead carrier and resume development ( Bovien, 1937 ; Richter, 1993 ). Typical examples are foun...

Ecology of Caenorhabditis species: 5. List of 23 Caenorhabditis species and their ecology
...this species phoresy is physiologically necessary to resume development ( Kiontke, 1997 ). Dauer juveniles do not actively disembar...


Ionotropic glutamate receptors: genetics, behavior and electrophysiology [HTML]
Ionotropic glutamate receptors: genetics, behavior and electrophysiology : 4. Electrophysiological analysis of iGluRs in C. elegans
... of the sensory neurons ( O'Hagan et al., 2005 ). Continued development of new stimulation techniques and their application to inte...

Ionotropic glutamate receptors: genetics, behavior and electrophysiology : 6. Mechanisms of iGluR localization, stabilization and function at the synapse
...d surface expression ( Stricker and Huganir, 2003 ). During development, the density of GLR-1 ::GFP puncta along the ventral cord r...

...ether, these findings suggested that UNC-43 activity during development is required to traffick GLR-1 from the cell body for the fo...


Trans-splicing and operons [HTML]
Trans-splicing and operons: 7. Signals on the polycistronic pre-mRNA for SL2 trans-splicing
...en found to immunoprecipitate the SL2 snRNP from C. elegans embryo extracts, and the region of SL2 RNA required for SL2 identi...


Electrophysiological recordings from the neuromuscular junction of C. elegans [HTML]
Electrophysiological recordings from the neuromuscular junction of C. elegans : 1. Introduction
... as well as additional insights derived from the Drosophila embryo dissection pioneered by Kendal Broadie ( Broadie, 1994 ). E...


Mechanosensation [HTML]
Mechanosensation : 1. Introduction
...t the molecules needed for the function, specification, and development of C. elegans MRNs than is known in other organisms. This c...

Mechanosensation : 5. Conclusions
...g complete lists of the genes needed for the specification, development and function of each MRN and improved understanding of how ...


Gene expression changes associated with aging in C. elegans [HTML]
Gene expression changes associated with aging in C. elegans
...5 USA This chapter is in WormBook section: > Post-embryonic development Publication History version 1 latest version Table of Conte...

Gene expression changes associated with aging in C. elegans : 4. What have we learned?
.... These genes are involved in processes such as metabolism, development, and detoxification and stress response, but intriguingly, ...


Maintenance of C. elegans [HTML]
Maintenance of C. elegans : 4. Culturing C. elegans on petri plates
...days so that you have a source of animals at every stage of development. 4.3. General stock maintenance tips C. elegans stocks...

Maintenance of C. elegans : 7. Freezing and recovery of C. elegans stocks
...successful freeze are using animals at the correct stage of development, the addition of glycerol to the freezing media, and a grad...


Natural variation and population genetics of Caenorhabditis elegans [HTML]
Natural variation and population genetics of Caenorhabditis elegans : 1. Molecular polymorphisms and population genetics
...iently, the maintenance of genes required for male-specific development and behavior seems to imply the existence of outcrossing in...

Natural variation and population genetics of Caenorhabditis elegans : 2. Phenotypic diversity
...r pheromone than some other strains ( Viney et al., 2003 ). Development. One of the few non-invariant cell division patterns in N2 ...


Transgenic solutions for the germline [HTML]
Transgenic solutions for the germline : 2. The germline challenge
...patterns ( Merritt et al., 2008 ). Second, most of germline development occurs in a syncytium, where germ cells in different develo...

Transgenic solutions for the germline : 4. Considerations when designing entry clones.
... 2008 ). Note that the rate of H2B turnover during germline development is not known and could in theory artificially extend or sho...


Gene expression changes associated with aging in C. elegans [HTML]
Gene expression changes associated with aging in C. elegans
...5 USA This chapter is in WormBook section: > Post-embryonic development View/Add Comments Publication History version 1 latest vers...

Gene expression changes associated with aging in C. elegans : 4. What have we learned?
.... These genes are involved in processes such as metabolism, development, and detoxification and stress response, but intriguingly, ...


Ethanol.pdf [preprint PDF]
Ethanol.pdf
...lated in Hawaii) than N2. These differences in the rate of development of acute tolerance were found to be due to allelic variati...

...sed seven times to the N2 strain, and selected for a rapid development of acute tolerance at each outcross, demonstrate more rapi...


Electrophysiological recordings from the pharynx [HTML]
Electrophysiological recordings from the pharynx : 2. Protocol 1. Extrapharyngeal recordings (EPGs)
...ple of an EPG is shown in Figure 1 . There is scope for new development here in the form of software for reliable automated analysi...

Electrophysiological recordings from the pharynx : 5. Acknowledgements
... Acknowledgements Irina Vinogradova contributed to the development of the patch clamp protocols. Enriq Claverol and Christophe...


Ethanol [HTML]
Ethanol : 6. npr-1 regulation of ethanol tolerance
...olated in Hawaii) than N2. These differences in the rate of development of acute tolerance were found to be due to allelic variatio...

...ssed seven times to the N2 strain, and selected for a rapid development of acute tolerance at each outcross, demonstrate more rapid...


Synaptic function [HTML]
Synaptic function : 4. Summary
...he vesicular ACh transporter UNC-17 , UNC-26 ). Through the development of behavioral tests imaging tools and electrophysiological ...

...ynaptic vesicle cycle outlined above. This will require the development of new optical tools to study vesicle dynamics, calcium han...


Signaling in the immune response [HTML]
Signaling in the immune response : 5. Role of Toll-like receptors
...d pathogenic S. marcescens , and is essential for embryonic development ( Pujol and Ewbank, 2003 ; Pujol et al., 2001 ). Activation...

...worms is different from that seen in the AWC neurons during development ( Figure 3 ). Significantly, the enhanced susceptibility to...


Pristionchus pacificus genomics: from genetics to genome sequence [HTML]
Pristionchus pacificus genomics: from genetics to genome sequence : 1. Introduction
...nts affecting several developmental processes such as vulva development, sex determination and gonad development ( Pires-daSilva and Sommer, 2004 ; Rudel et al., 2005 ; Som...


RNA in situ hybridization of dissected gonads [HTML]
RNA in situ hybridization of dissected gonads : 1. Introduction
...rtant step in understanding gene function during C. elegans development. The expression pattern of a gene can be examined in severa...


SNPs: Introduction and two-point mapping [HTML]
SNPs: Introduction and two-point mapping : 2. The SNP databases
...s somewhat more preliminary and the site is currently under development. However, since this database has been assembled using the ...


Karyotype, ploidy, and gene dosage [HTML]
Karyotype, ploidy, and gene dosage: 8. Dosage sensitive genes
...ulated by translational control, in order to achieve proper development of the hermaphrodite germ line (see Sex-determination in th...


Genomic classification of protein-coding gene families [HTML]
Genomic classification of protein-coding gene families: 3. Classifying proteins
...h aligned families ( Durbin et al., 1999 ). This led to the development of the HMMER search software ( Eddy, 2005 ) and its use to ...


Genetic mapping and manipulation: Chapter 6-Mapping with deficiencies and duplications [HTML]
Genetic mapping and manipulation: Chapter 6-Mapping with deficiencies and duplications : 1. Basics
...genes usually includes some that are necessary during early development. One of the traditional strengths of Df mapping is that it ...


Interactions with microbial pathogens [HTML]
Interactions with microbial pathogens : 3. Bacterial infections of the cuticle
...e. Mutations in let-60 , which activates the ERK pathway in development, did not produce Bus, suggesting a different means of activ...


Genetic mapping and manipulation: Chapter 6-Mapping with deficiencies and duplications [HTML]
Genetic mapping and manipulation: Chapter 6-Mapping with deficiencies and duplications : 1. Basics
...genes usually includes some that are necessary during early development. One of the traditional strengths of Df mapping is that it ...


Egg-laying [HTML]
Egg-laying: 2. Functional roles of egg-laying circuitry components
...te, and many egg-laying defective mutants have abnormal HSN development or function ( Trent et al., 1983 ; Desai and Horvitz, 1989 ...


Heterologous expression of C. elegans ion channels in Xenopus oocytes [HTML]
Heterologous expression of C. elegans ion channels in Xenopus oocytes
... of C. elegans, a model system used for many years to study development and a plethora of biological processes mainly employing gen...


Potassium channels in C. elegans [HTML]
Potassium channels in C. elegans : 5. Usefulness of the information in C. elegans
...s as chronic pain. Naturally such uses have to wait for the development of tissue-specific promoters and viral or “gene-gun&#...


Web resources for C. elegans studies [HTML]
Web resources for C. elegans studies : 2. Portal and knowledge environment
...ev.wormbase.org/BioMart/martview > Presently still under development, WormMart is an advanced query tool which allows users cust...

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