Gametogenesis
The series of transformations that results
in the formation of mature gametes is
called gametogenesis. Although the
same essential processes are involved in
the maturation of both sperm and eggs,
there are some important differences.
Gametogenesis in testes is called
spermatogenesis, and in ovaries,
oogenesis.
Spermatogenesis
The walls of the seminiferous tubules
contain differentiating germ cells
arranged in a stratified layer five to eight
cells deep (Figure 7-7). Germ cells
develop in close contact with large
sustentacular (Sertoli)
cells, which extend
from the periphery of the seminiferous
tubules to the lumen and provide nourishment
during germ cell development
and differentiation (Figure 7-8). The outermost
layers contain spermatogonia,
diploid cells that have increased in number
by ordinary mitosis. Each
spermatogonium increases in size and becomes a
primary spermatocyte. Each primary
spermatocyte then undergoes the first
meiotic division, as described previously,
to become two
secondary spermatocytes.
Each secondary spermatocyte
enters the second meiotic division
without the intervention of a resting
period. In the two steps of meiosis
each primary spermatocyte gives rise
to four spermatids, each containing
the haploid number (23 in humans) of
chromosomes. A spermatid may contain
all chromosomes
that the male
inherited from his mother, those he
inherited from his father, or most
likely, a combination of his parents’
chromosomes. Without further divisions
the spermatids are transformed
into mature
spermatozoa or (
sperm)
(Figure 7-8).
Modifications include
great reduction of cytoplasm, condensation
of the nucleus into a head, formation
of a middle piece containing
mitochondria, and a whiplike, flagellar
tail for locomotion (Figure 7-8, 7-9).
The head consists of a nucleus containing
the chromosomes for heredity
and an
acrosome, a distinctive feature
of nearly all the metazoa (exceptions
are teleost fishes and certain invertebrates).
In many species, both invertebrate
and vertebrate, the acrosome
contains lysins that serve to clear an
entrance through the layers that surround
the egg. In mammals at least,
one of the lysins is the enzyme
hyaluronidase, which allows the sperm
to penetrate the follicular cells surrounding
the egg. A striking feature of
many invertebrate spermatozoa is the
acrosome filament, an extension of
varying length in different species that
projects suddenly from the sperm head
when the latter first contacts the surface
of the egg. The fusion of the egg
and sperm plasma membranes is the
initial event of fertilization (See Contact
and Recognition between Egg and
Sperm).
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Figure 7-7 Section of a seminiferous
tubule containing male
germ cells.
More than 200 long, highly coiled
seminiferous tubules are packed in
each human
testis. This scanning
electron micrograph reveals,
in the
tubule’s central cavity, numerous
tails of
mature spermatozoa that
have differentiated from
germ cells in
the periphery of the tubule. (×525) |
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Figure 7-8 Spermatogenesis. Section of seminiferous tubule showing
spermatogenesis. Germ cells develop within the recesses of large
sustentacular (or Sertoli) cells,
that extend from the periphery of
seminiferous tubules to their lumen, and that provide nourishment to the
germ cells. Stem germ cells from which the
sperm differentiate are the
spermatogonia, diploid cells located peripherally in the tubule. These divide
by mitosis to produce either more spermatogonia or
primary
spermatocytes. Meiosis begins when the primary spermatocytes divide to
produce haploid secondary spermatocytes with double-stranded
chromosomes. The second meiotic division forms four haploid spermatids
with single-stranded chromosomes. As the sperm develop they are
gradually
pushed toward the lumen of the seminiferous tubule. |
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The total length of a human sperm
is 50 to 70 µm. Some toads have sperm
that exceed 2 mm (2000 µm) in length
(Figure 7-9) and are easily visible to
the unaided eye. Most sperm, however,
are microscopic in size. In all sexually
reproducing animals the number
of sperm in males is far greater than
the number of eggs in corresponding
females. The number of eggs produced
is related to the chances of the young
to hatch and reach maturity.
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Figure 7-9 Types of vertebrate and invertebrate sperm. |