Z
Z Chromosome
D W Burt
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.1404
Avian and mammalian sex chromosomes evolved
independently (Fridolfsson et al., 1998; Nanda et al.,
1999, 2000) and should therefore have fundamentally
different sex-determining genes. The sex chromo-
somes in birds are designated Z and W: the female is
the heteromorphic (ZW) and the male the homo-
morphic (ZZ) sex. The average avian Z chromosome
is a medium-sized macrochromosome. It is not clear
in birds whether the Z or W chromosome determines
sex.
References
Fridolfsson A-K, Cheng H, Copeland NG et al. (1998) Evolution
of the avian sex chromosomes from an ancestral pair of
autosomes. Proceedings of the National Academy of Sciences,
USA 95: 8147±8152.
Nanda I, Shan Z, Schartl M et al. (1999) 300 million years of
conserved synteny between chicken Z and human chromo-
some 9. Nature Genetics 21: 258 ±259.
Nanda I, Zend-Ajusch E, Shan Z et al. (2000) Conserved syn-
teny between the chicken Z sex chromosome and human
chromosome 9 includes the male regulatory gene DMRT1:
a comparative (re)view on avian sex determination.
Cytogenetics and Cell Genetics 89: 67±78.
See also: Sex Determination, Human;
W Chromosome
Z DNA
J H Miller
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.1405
A zig-zag-like structure of the DNA chain that is
observed in GC-rich segments of DNA which form
left-handed helices.
Zinc Finger Proteins
See: DNA-Binding Proteins
Zoo Blot
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.2073
A zoo blot is a technique using Southern blotting to
evaluate the ability of a DNA probe from one species
to hybridize with genomic DNA from a variety of
other species.
See also: Southern Blotting
Zygote
P M Wassarman
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.1407
The zygote, or fertilized egg, is a single cell produced
by fusion of female and male germ cells, that is, the
unfertilized egg and sperm, respectively. Since germ
cells undergo meiotic divisions to a haploid state (n)
during oogenesis and spermatogenesis, fusion of the
unfertilized egg and sperm (fertilization) restores a
diploid (2n) number of chromosomes to the zygote.
In mammals, the second meiotic division of the egg,
with separation of chromatids, occurs shortly after
fusion with sperm. At an appropriate time after fertil-
ization, the zygote begins to divide mitotically, even-
tually giving rise to a multicellular organism that
exhibits all of the characteristics of the species.
Nuclei contributed to the zygote by the unfertil-
ized egg and sperm are called female and male pronu-
clei, respectively. In mice, the female pronucleus forms
at *7.5 h and the male pronucleus at *5.5 h after
fusion of the unfertilized egg and sperm. The two
pronuclei must come together near the center of the
zygote and form a single diploid nucleus. The timing
of nuclear formation varies greatly from one species to
another; for example, it takes  1 h in sea urchins and
2166 Z y g o t i c L e th a l G en e
 12 h in mice. In fact, in mice, pronuclei approach each
other, but do not actually fuse to become a diploid
nucleus. Rather, pronuclear membranes disappear and
chromosomes assemble on a spindle. DNA replication
occurs *14.5 h after fertilization, as the pronuclei
migrate toward the center of the zygote. In mice, the
first cleavage division occurs at *20 h after fertil-
ization, when chromosomes are assembled on a spindle.
In many animals, sperm contribute a centriole to
the zygote and this organelle helps to organize the
first mitotic spindle on which the chromosomes are
arranged. In this respect, the sperm centriole acts
as a microtuble-organizing center in the zygote. On
the other hand, sperm contribute very few of the
large number of mitochondria found in the zygote
( 0.01%), ensuring that mitochondrial DNA is
maternally inherited.
The zygote is inactive with respect to nascent
transcription of genomic DNA, although translation
of maternal transcripts takes place. The onset of tran-
scription is delayed until after the first cleavage division
in mammals and after the first 12 cleavage divisions in
some nonmammals. Presumably, this period of tran-
scriptional inactivity exhibited by the zygote provides
time to remodel parental chromosomes.
In mammals, genomes derived from the unfertil-
ized egg and sperm appear not to be equivalent. In
some cases, only the maternally derived allele of a
particular gene is active, whereas, in others, only the
paternally derived allele is active (`genetic imprint-
ing'). Some of these genes are absolutely essential for
normal development. Apparently, as a result of this
nonequivalence of pronuclei, parthenogenetic (bima-
ternal), gynogenetic (bimaternal), and androgenetic
(bipaternal) mammalian zygotes cannot give rise to
normal fetuses and live births.
Further Reading
Gilbert SF (1997) Developmental Biology, 5th edn, p. 918.
Sunderland, MA: Sinauer Associates.
Hogan B, Beddington R, Costantini F and Lacy E. (1994)
Manipulating the Mouse Embryo, 2nd edn, p. 497. Plainview,
NY: Cold Spring Harbor Laboratory Press.
See also: Fertilization
Zygotic Lethal Gene
R K Herman
Copyright ß 2001 Academic Press
doi: 10.1006/rwgn.2001.1408
A zygotic lethal gene is a gene that leads invariably or
almost invariably to the death of an organism prior to
the reproductive stage. `Zygotic' in this case refers
to the organism that develops from a single-celled
diploid zygote. For some zygotic lethal genes, a rare
individual carrying the gene in a dose that is normally
lethal survives to the reproductive stage. Such rare
survivors are called `escapers.' Zygotic lethal genes
are to be distinguished from the following: gametic
or haplophasic lethal genes, which exert their effects in
haploid gametes; from sterile genes, which allow their
bearers to reach reproductive maturity but render
them sterile; and from maternal-effect or paternal-
effect lethal genes, which kill the progeny of affected
individuals.
Many zygotic lethal genes result in developmental
arrest and death at a particular stage during develop-
ment, referred to as the lethal phase. Zygotic lethal
genes may thus be classified as embryonic or postem-
bryonic, depending on their lethal phase. Examples of
postembryonic zygotic lethal genes in insects include
larval, pupal and early adult lethals.
A dominant zygotic lethal gene is lethal when pres-
ent as a single copy per cell, even when a wild-type
allele of the same gene is also present in the diploid
cells. Such lethal genes are rarely studied because they
cannot be propagated by breeding. Recessive zygotic
lethal genes are lethal only when they are present in
the homozygous or hemizygous condition. Indivi-
duals that are heterozygous for the lethal gene are
viable because the wild-type allele is dominant to the
lethal allele. A recessive lethal gene can be maintained
and propagated in a heterozygous stock. One-fourth
of the progeny produced from the mating of hetero-
zygous lethal parents are expected to be homozygous
for the lethal gene and exhibit the lethal phenotype.
Two-thirds of the surviving progeny are expected to
be heterozygous for the lethal gene and can be used
to propagate the lethal gene. In rare cases, a recessive
zygotic lethal gene may confer a visible phenotype
dominantly, in which case individuals heterozygous
for the lethal can be readily identified. For most lethal
genes, however, the heterozygous lethal organisms are
indistinguishable from the homozygous wild-type
organisms, unless a closely linked marker gene con-
ferring a visible phenotype is used to track either the
lethal-bearing chromosome or its non-lethal homo-
log. Chromosomal rearrangements are often used to
suppress recombination between the lethal gene and
the visible tag.
X-linked zygotic lethal genes have long (since
1912) been recognized in the fruit fly Drosophila
melanogaster because the progeny of a heterozygous
mother mated to a wild-type male exhibit an altered
sex ratio: half of the sons are hemizygous for the
zygotic lethal gene and inviable, whereas all of the
daughters receive a dominant wild-type allele from