Worm Breeder's Gazette 11(2): 110

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A Survey of the Male Phenotypes of Some egl Mutants

Andrew Chisholm

Figure 1

The purpose of this survey was to see if any of the egl mutants had 
abnormal male phenotypes, and to find out what proportion of a random 
sample of mutations isolated on the basis of a sex-limited phenotype 
were wholly sex-specific in their effects.
A number of egl mutations turned out to cause abnormal male 
phenotypes, described below.  Out of a total of 42 egl mutations 
inspected, only 12 appeared to have no phenotype outside of the 
egglaying system, and may define genes with roles confined to the 
hermaphrodite sex.  However, for most of the loci analyzed, the null 
phenotype was undefined, so any conclusions regarding sex-specificity 
are tentative.  Some male phenotypes had been described previously (
Trent et al., 1983; Desai and Horvitz, 1989); my observations confirm 
theirs.
egl mutations of 26 loci had no discernible male phenotype: males 
appear wild-type and mate with ME of 2/3.  Mutations of 6 loci (egl-3, 
egl-19, egl-33 and egl-39 ) 
had male mating defects which could be attributed to a non-sex-
specific phenotype such as Unc; egl-8 and egl-30 males also have 
slight morphological defects.  Mutations at 11 loci caused a male 
mating defect, apparently the result of morphological defects.  egl-5 
and egl-27 have grossly abnormal male tails as a result of abnormal 
tail lineages.  The egl-31 mutation causes abnormal M lineages in both 
sexes.  egl-8, egl-20, egl-34 
and egl-45 males all have variable morphogenetic defects in the tail.  
egl-28 and egl-32 males have gonad defects.  egl-2 males have abnormal 
bursal morphology, but this may result from a muscular or neural 
defect.
egl mutations of 3 loci caused a male mating defect for no apparent 
reason.  egl-4, pear 
morphologically normal but mate very poorly.  It is possible that 
there is some subtle defect in male mating which I have not noticed, 
or possibly a defect in male sperm function.
egl-41 males have slightly reduced mating efficiency, presumably the 
result of the weak feminization of egl-41 XOs.
Abnormal male 
phenotypes
egl-2 Males are well-coordinated and can back and turn, but cannot 
locate the vulva on hermaphrodites.  The structures of the tail are 
all present and morphologically normal, but the cloacal structures (
including hook, spicules and gubernaculum) are permanently protruding 
in the position normally only seen during copulation.  The hook and 
its sensillum are almost essential for vulval location (WBG 7,2) and 
it is possible that in these priapic worms the hook cannot operate 
properly.  n693 is semidominant for this phenotype: n693/+ males mate 
reasonably well, but a small fraction have extruded cloacas.  In a 
preliminary experiment the mating defect was rescued by exogenous 
imipramine.
egl-8 Males are Unc and slightly thin, and about one third have 
morphological defects in the fan: rays may be missing and the 
posterior fan may bulge out ventrally.  Some males are quite severely 
affected.
egl-14 Males could back and turn in mating, but appeared to locate 
the vulva very poorly.
egl-18 Males have variable tail fan morphological abnormalities: 
rays may be blobby, misshapen or lost altogether.  There is also a low 
penetrance Vab phenotype, both sexes.
egl-20 At 20 C and 25 C n585 males have slightly abnormal tail 
morphogenesis (blobby rays, small fan) and variable defects in their 
sex muscles.  Often the more anterior diagonal muscles appear to be 
missing or misplaced; sometimes ectopic diagonal-like muscles can be 
seen attaching in the midbody or more anteriorly.  Some males have the 
crumpled spicules characteristic of sex muscle defects, although the 
other sex muscles appear to be present.  In some animals the dorsal 
coelomocyte could not be seen.  Occasionally the first ray appeared 
not to have migrated posteriorly, causing the alae to have a gap or 
end anteriorly to normal.
egl-27  There are several interesting phenotypes in egl-27 mutants; 
I hope to describe the male lineage defects in detail at some time in 
the future.
egl-28  Male tail morphology is mostly wild-type, although a small 
fraction of males have minor morphogenetic defects (such as a small 
fan).  In some egl-28 males the germline has an unusual appearance: 
some sperm appear to be made initially, but the germ-line more distal 
to the sperm has a dough-like appearance and does not appear to be 
differentiating into sperm.  This phenotype has low penetrance at 20 C 
and 25 C.
egl-32  Male tail morphology was wild-type, but variable 
abnormalities in male somatic gonads and germline were seen.  These 
defects are incompletely penetrant.  At 20 C, 4/67 males had abortive 
somatic gonad development, in which the gonad ended up as a 
structureless mass instead of a reflexed tube.  Some sperm could be 
seen differentiating in the blob.  15/67 had germlines of abnormal 
appearance.  At 25 C, 1/40 had the severe gonad defect and 12/40 had 
germline abnormalities.  In 2 of the 12, gametes with the morphology 
of oocytes were seen.  Neither the gonad nor the germline phenotype 
was seen in 79 egl-32 males grown at 15 C.  These phenotypes are 
reminiscent of those described for fog-1.  Alleles of fog-1 can cause 
feminization of the germline in both sexes, and have poorly understood 
effects on the male somatic gonad at low penetrance (T.  Doniach, pc).
egl-32 may have functions similar to those of fog-1, n155 being a 
weak allele with a minor effect on the hermaphrodite germline giving 
the Egl phenotype.  Alternatively, since egl-32 and fog-1 are linked, 
there may be a weak allele of fog-1 in the strain; I have not tried to 
check this.
egl-34 Males are variably morpho-mab, some having blobby rays and 
some a misshapen ventral bursa: the hook can appear displaced 
anteriorly from the cloaca.  Variable sex muscle defects also occur: 
some males have crumpled spicules, and occasionally some diagonal 
muscles are missing.
egl-35  At 25 C males appear variably morpho-mab (misplaced rays, 
swollen bursa) and occasionally have crumpled spicules.  This defect 
is weaker at 20 C.
egl-45  Males are variably morpho-mab: some appear almost WT, while 
some have posterior bulges in the fan similar to those seen in mab-9 
males.  Some have crumpled or missing spicules and very variable sex 
muscle defects.
[See Figure 1]

Figure 1