WormMine: LifeStage WBls:0000021

• Definition: The C. elegans life stage spanning 620-800min(hatch) after first cleavage at 20 Centigrade. Cell number remains at ~560 cells, with some new cells generated and some cells go through programmed cell death. A stage after elongation is over. The last stage of embryogenesis. Also called pre-hatched embryo, late embryo or morphogenetic embryo.
• Primary Identifier: WBls:0000021
• Public Name: fully-elongated embryo Ce

555 Anatomy Terms

Definition	Name	Synonym	Primary Identifier
Pharyngeal sensory-motorneurons	m3R	lineage name: ABaraappppp	WBbt:0004469
Ring interneuron	URXL	lineage name: ABplaaaaappp	WBbt:0004928
Ring interneuron	URXR	lineage name: ABarpapaappp	WBbt:0004926
Pharyngeal sensory-motorneurons	m3L	lineage name: ABaraapappp	WBbt:0004471
Neuron class of one interneuron, projects along ventral cord to ring.	PVT	lineage name: ABplpappppa	WBbt:0004070
gland cell of the secretory-excretory system, sends processes to ring, opens into excretory duct.	excretory gland cell	exc gl	WBbt:0005776
Herm. specific motor neurons (die in male embryo), innervate vulval muscles, serotonergic	HSNL	lineage name: ABplapppappa	WBbt:0004758
Herm. specific motor neurons (die in male embryo), innervate vulval muscles, serotonergic	HSNR	lineage name: ABprapppappa	WBbt:0004757
Amphid interneuron	AIYL	lineage name: ABplpapaaap	WBbt:0003963
Amphid interneuron	AIYR	lineage name: ABprpapaaap	WBbt:0003961
one member of Amphid neurons, single ciliated endings, probably chemo-sensory; project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil	ASEL	lineage name: ABalppppppaa	WBbt:0003904
one member of Amphid neurons, single ciliated endings, probably chemo-sensory; project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil	ASER	lineage name: ABpraaapppaa	WBbt:0003903
Neuron, ciliated ending in head, no supporting cells, associated with ILso	BAGL	lineage name: ABalppappap	WBbt:0003824
Neuron class of one pharyngeal interneuron, posterior sensory.	I5 neuron	I5	WBbt:0004740
one member of Amphid neurons, single ciliated endings, probably chemo-sensory; project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil	ASGL	lineage name: ABplaapapap	WBbt:0003892
one member of Amphid neurons, single ciliated endings, probably chemo-sensory; project into ring via commissure from ventral ganglion, make diverse synaptic connections in ring neuropil	ASGR	lineage name: ABpraapapap	WBbt:0003891
Neuron class of one pharyngeal motorneuron.	M5 neuron	lineage name: MSpaaapap	WBbt:0004465
Neuron, ciliated ending in head, no supporting cells, associated with ILso	BAGR	lineage name: ABarappppap	WBbt:0003823
H-shaped cell associated with the excretory system, largest cell in C. elegans.	excretory cell	excretory canal cell	WBbt:0005812
Inner labial neuron	IL1VR	lineage name: ABarapppppaa	WBbt:0004547
Inner labial neuron	IL1DR	lineage name: ABalappppaaa	WBbt:0004737
Inner labial neuron	IL1L	lineage name: ABalapaappaa	WBbt:0004553
Inner labial neuron	IL1R	lineage name: ABalaappppaa	WBbt:0004551
Inner labial neuron	IL1VL	lineage name: ABalppapppaa	WBbt:0004549
Inner labial neuron	IL1DL	lineage name: ABalapappaaa	WBbt:0004738
Amphid socket cell, right	AMsoR	lineage name: ABprpaapapa	WBbt:0003929
Amphid neuron, prob. chemosensory	ADFL	lineage name: ABalpppppaa	WBbt:0004003
Amphid neuron, prob. chemosensory	ADFR	lineage name: ABpraaappaa	WBbt:0003999
Ring interneuron	AIML	lineage name: ABplpaapppa	WBbt:0003983
Ring interneuron	AIMR	lineage name: ABprpaapppa	WBbt:0003969

2 Contained In

Remark	Definition	Other Name	Public Name	Primary Identifier
	The whole period of embryogenesis in the nematode Caenorhabditis elegans, from the formation of an egg until hatching.		embryo Ce	WBls:0000003
	A C. elegans life stage that occurs during the fully-elongating embryo life stage		C. elegans life stage occurring during fully-elongated embryo	WBls:0000811

0 Expression Clusters

819 Expression Patterns

• Primary Identifier: Expr8361
• Remark: Picture: Figure 1.
• Pattern: GFP expression initiated in the early gastrula. Robust expression of Prncs-1::GFP was observed in the midgut (E cell lineage) starting at the 28-cell stage and continuing into adulthood. By the comma stage, fluorescence was also visible in the embryo periphery in cells that give rise to hypodermis. In L1 larva and subsequent stages, strong expression of GFP was seen in hypodermal cells, including Hyp 7 syncytium and head and tail hypodermis. The expression pattern was identical in hermaphrodites and males, but adult hermaphrodites displayed fluorescence in vulval epithelium. Expression was absent in seam cells, nervous system, and pharynx. The Prncs-1::GFP reporter showed increased expression during starvation. Although fluorescence intensity was enhanced under starved conditions, the spatial expression pattern was unchanged. Expression of the Prncs-1::GFP transgene was also enhanced in males. An ~2.5-fold increase in rncs-1 expression in total RNA prepared from wild-type, well fed males, compared with hermaphrodites.

• Primary Identifier: Expr4883
• Remark: Picture: Fig. 4A.
• Pattern: Low GFP signals were detected exclusively in the intestinal cells of late embryos, L1L4, and adult hermaphrodites.

• Primary Identifier: Expr4836
• Remark: Picture: Figs. 4A-D.
• Pattern: In hermaphrodites, the expression of bro-1 was restricted to seam cells. Its expression was first detected at bean-stage embryos and persisted throughout the developmental stages. In the male tail, bro-1 was also expressed in the ray precursor cells.
• Subcellular Localization: GFP::BRO-1 was localized to both the cytoplasm and the nucleus.

• Primary Identifier: Expr4837
• Remark: Picture: Figure 5.
• Pattern: Fluorescence started to be visible in two cells of young embryos at around the 64 AB cell stage. Towards the end of gastrulation expression was visible in about 40 cells throughout the embryo including neuronal precursors, ventral hypodermal cells, and pharyngeal precursor cells. At the 1 to 2 fold stages fluorescence was observed in IL1 neurons (the identity was determined post-embryonically), the nine buccal epidermal cells, and additional cells in the head, most likely arcade cells. Transient expression was also observed in embryonic motoneurons (no longer visible in 3 fold stage embryos) and in a few apoptotic cells in the head. Based on their position they could be the sister cells of some of the IL1 neurons, which are known to undergo programmed cell death at this developmental stage. At the 3 fold stage expression was restricted to the buccal epidermal cells, most of the arcade cells (3 anterior and the DL and DR posterior arcade cells), and the six IL1 neurons. The two lateral IL1 neurons expressed the marker only weakly also in the L1 larval stage (but not later during development), whereas the dorsal and ventral IL1 neurons expressed GFP strongly throughout all larval stages and in the adults. Starting from the L1 larval stage expression could also be observed in the posterior cells of the gut. Starting from the L2 stage, when gonad development and migration begins, fluorescence became also visible in the distal tip cells of the gonad.

• Primary Identifier: Expr4829
• Remark: Picture: Fig. 6A, 6B. Reporter gene fusion type not specified.
• Pattern: Exclusively expressed throughout the nervous system in C. elegans. F25B3.3::gfp is a postmitotic pan-neuronal marker, i.e. its onset of expression is observed after the terminal division of neurons (around 450 minutes of embryonic development).

• Primary Identifier: Expr4813
• Remark: Picture: Figure 7, C and D.
• Pattern: Expression was observed throughout development, starting at midembryogenesis. VHA-5 was also detected at the lumen of the vulva and rectum. In addition, authors found VHA-5 expressed in the sheath cells associated with head and tail sensory organs. Three-dimensional reconstructions showed that VHA-5 and RDY-2 formed a sixfold symmetrical pattern, which includes a larger spot that presumably corresponds to the amphid.
• Subcellular Localization: In the amphid sheath cell, VHA-5 was found in the most distal part of the cell lining the sheath pocket, which can be equated to its apical side.

• Primary Identifier: Expr4693
• Remark: nsy-5 = T16H5.1.
• Pattern: A GFP reporter transgene with 5.8 kb of the nsy-5 promoter was expressed exclusively in sensory neurons and interneurons in the head and tail. The neurons that expressed nsy-5::GFP included AWC, ASH, AFD, ASI, ADL, ASK, BAG, AWB, and ADF (head sensory neurons); ADA, AIZ, RIC, AIY, and AIM (head interneurons); PHA and PHB (tail sensory neurons); and PVC and PVQ (tail interneurons). Expression began about halfway through embryogenesis, was strongest in late embryogenesis and the L1 larval stage, and faded thereafter. Adults maintained weak expression in several neurons, including ASH but not AWC.

• Primary Identifier: Expr4788
• Pattern: A full-length glt-3-gfp fusion (later shown to be capable of rescuing the phenotypes of a glt-3 mutant strain) is expressed at increasing levels from late embryogenesis to adulthood throughout the body-spanning excretory canal cell.
• Subcellular Localization: The expression of GLT-3::GFP appears localized to the abluminal (basolateral) side.

• Primary Identifier: Expr4767
• Pattern: Young embryos did not present a GFP signal, but expression was observed in late embryogenesis and at all stages of postnatal development: eggs, L1, L2, L3, L4, and adults. Adult animals showed stronger expression than did larval stages and eggs; this was also proved by RT-polymerase chain reaction (RT-PCR), suggesting potential developmental dynamics in atx-3 function. Both transgenic strains had a generalized expression pattern, with a strong signal in the spermatheca and vulval muscle . High fluorescence was observed in neuronal dorsal and ventral cord and neurons of the head and tail. Expression was also observed in the hypoderm, body muscles, and coelomocytes.

• Primary Identifier: Expr4735
• Remark: Reporter gene fusion type not specified.
• Pattern: Expression began at the comma stage and continued from the L1 to adult stages in body wall muscles. The tni-1::lacZ animals showed only weak expression in the body wall muscles.

• Primary Identifier: Expr4736
• Remark: Reporter gene fusion type not specified.
• Pattern: Expression began at the comma stage and continued from the L1 to adult stages in body wall muscles. The tni-2::lacZ construct gave strong expression in all the body wall muscles.

• Primary Identifier: Expr4737
• Remark: Reporter gene fusion type not specified.
• Pattern: Expression began at the comma stage and continued from the L1 to adult stages in body wall muscles. The tni-3::lacZ construct was expressed in all the body wall muscles from the L1 to adult stages. In the adult stage, tni-3::lacZ expression in body wall muscles became weaker with time although strong expression persisted in the head and vulval muscles.

• Primary Identifier: Expr4738
• Remark: Reporter gene fusion type not specified.
• Pattern: tni-4::lacZ was expressed only in the pharynx from the 2-fold to adult stages.

• Primary Identifier: Expr4986
• Remark: Picture: Figure 5B.
• Pattern: Consistent with the ubiquitous expression of sqt-3 throughout the life cycle, the R584 antiserum intensely stains the hypodermal cells and cuticles of all stages. Reactive antigens are first detected in wild-type comma-stage embryos within the hypodermal cells. The signal localizes perinuclearly, presumably in association with the secretory pathway. At the threefold stage, coincident with cuticle secretion, the signal becomes progressively extracellular: by the late pretzel stage, all antigen is detected in alignment with the annular ridges of the embryonic cuticle.

• Primary Identifier: Expr4954
• Remark: Picture: Fig. 2.
• Pattern: In larvae and adults, the circumferential filamentous pattern did not persist, but expression was seen in a subset of head and tail socket cells, the vulva, more faintly the uterus, and the rectum.
• Subcellular Localization: GFP expression was observed in the epidermis from the 1.2-fold stage of elongation to the end of embryogenesis. The GFP formed a circumferential filamentous pattern that was spatially and temporally strikingly reminiscent of the circumferential actin microfilament pattern. Indeed, actin staining in a strain expressing the RGA-2::GFP construct showed that the two networks coincide.

• Primary Identifier: Expr4956
• Remark: Picture: Fig. 5, Fig 6.
• Pattern: NAB-1::GFP expression is restricted to epithelia and neurons. The earliest expression was observed in the hypodermis of 2-fold-stage early embryos. Immediately prior to hatching, this expression became restricted to the epithelial excretory canal and the nervous system, including the central nervous system and the motoneurons (dorsal and ventral nerve cords. In L3 and L4 larvae, NAB-1::GFP also localized transiently at the membranes of the developing vulva epithelia. Also expressed in distal tip cell (pers. comm. from Wesley Hung 11-17-07.)
• Subcellular Localization: NAB-1::GFP puncta partially co-localized with the synaptic-vesicle protein SNT-1 and the active-zone protein UNC-10, suggesting that NAB-1 is present in presynaptic regions that are associated with vesicle pools and active zones. Similar to NAB-1, SAD-1 also showed co-localization with SNT-1. NAB-1::GFP and SAD-1 also showed partial co-localization, where each NAB-1::GFP punctum was associated with SAD-1 staining.

• Primary Identifier: Expr4940
• Pattern: strong bwm, vul, mu int, occaisonal weak pharyngeal, spermatheca, faint VNC, head neurons; embryonic bwm starts at 3-fold and very strong in L1s.

• Primary Identifier: Expr4941
• Pattern: strong bwm in 3-fold embryos; no other muscle seen in larvae or adults; saw faint bwm in 1.5 fold embryo but no earlier.

• Primary Identifier: Expr4942
• Pattern: strong bwm, vul, anal dep, mu int; strain A shows embryonic bwm starting at 1.5 fold and strong at 3-fold

• Primary Identifier: Expr4943
• Pattern: strong 3-fold emb bwm that fades and becomes mosaic in late L's and adults, vul, anal dep, spinchter, mu int; seam cells in one line at least

• Primary Identifier: Expr4944
• Pattern: strong bwm, vulval muscle and anal depressor, strong bwm at 3-fold into L1s, earliest exp is faint bwm in 2-fold seen in line B.

• Primary Identifier: Expr4931
• Pattern: strong bwm, pharyngeal, vulval, and anal depressor; a bit mosaic; starts at 3-fold?

• Primary Identifier: Expr4926
• Pattern: strong pan neuronal; spermatheca; faint vulval mu; rare and faint bwm in adult; neuronal starts at bean with strong at 3-fold.

• Primary Identifier: Expr4924
• Pattern: strong GFP expression in BWM, anal muscle and vulval muscles. GFP visible in late embryo.

• Primary Identifier: Expr4915
• Pattern: Strong but mosaic bwm; spermatheca; head neurons; 3-fold embryo strong bwm, no earlier embryonic seen.

• Primary Identifier: Expr4916
• Pattern: Strong and mosaic bwm expression beginning at 3-fold; no other muscle types seen.

• Primary Identifier: Expr4917
• Pattern: Moderate bwm, mosaic, tail hyps, earlies bwm is comma with 1-2 cells +, by 3-fold fairly strong in many bwm; early gut at 12E, strong gut at 1.5 fold then fades to cytoplasmic and gone.

• Primary Identifier: Expr4919
• Pattern: Strong pharyngeal, vulval, bwm, anal dep, mu int; 3-fold earliest embryonic.

• Primary Identifier: Expr4914
• Pattern: Strong GFP expression in BWM, anal muscle and vulval muscles. GFP visible in late embryo.

• Primary Identifier: Expr4393
• Pattern: The GFP signal was first detected from the 3-fold stage of embryogenesis during development. At the 3-fold stage, GFP expression was observed in hypodermal and intestinal tissues. During larval and adult stages, GFP was strongly detected in the hypodermis and intestine as well as the excretory cells.

1 Followed By

Remark	Definition	Other Name	Public Name	Primary Identifier
	The stage that begins right after hatching and before development proceeds after feeding. A newly hatched L1 larva contains 558 cells.		newly hatched L1 larval stage Ce	WBls:0000801

1 Preceded By

Remark	Definition	Other Name	Public Name	Primary Identifier
	The C. elegans life stage spanning 350-620min after first cleavage at 20 Centigrade. Cell number remains at ~560 cells, with some new cells generated and some cells go through programmed cell death. The stage that embryo starts elongation until elongation is over.		elongating embryo Ce	WBls:0000015

6 Sub Stages

Remark	Definition	Other Name	Public Name	Primary Identifier
	The time period encompassing the 640th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		640 min post first-cleavage Ce	WBls:0000695
	The time period encompassing the 650th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		650 min post first-cleavage Ce	WBls:0000696
	The time period encompassing the 660th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		660 min post first-cleavage Ce	WBls:0000697
	The time period encompassing the 720th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		720 min post first-cleavage Ce	WBls:0000698
	The time period encompassing the 770th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		770 min post first-cleavage Ce	WBls:0000699
	The time period encompassing the 780th minute post first cleavage at 20 Centigrade with respect to Sulston's lineage tree.		780 min post first-cleavage Ce	WBls:0000700