sperm oogenesis
TRANSCRIPT
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Ploidy: Number of sets of
chromosomes in a cell
Haploid (n)-- one set chromosomes
Diploid (2n)-- two sets chromosomes
Eggs and sperm are haploid (n)
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Developmental stages of
spermatogenesis
In the course of spermatogenesis the germ cellsmovetowards the lumen as they mature.
A-spermatogonium
-spermatogonium
!rimary spermatocyte (" spermatocyte order I)
#econdary spermatocyte (" spermatocyte order II)
#permatid #perm cell (" spermato$oon)
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%he spermatogenesis can &e su&di'ided into
two successive sections
%he first comprises the cells from thespermatogonium up to and including the
secondary spermatocyte and is termed
spermatocytogenesis. %he second one comprises the
differentiationmaturation of the sperm cell*
starting with the spermatid phase and istermed spermiogenesis (or
spermiohistogenesis).
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%he appro+imate 64 daycycle of the
spermatogenesis can &e su&di'ided into four
phases that last differing lengths of time
,itosis of the spermatogonia-- days--- /p to the
primary spermatocytes
0irst meiosis-- 21 days-- 0or the di'ision of the primary
spermatocytes to form secondary spermatocytes
#econd meiosis-- A few hours-- 0or the spermatids#permiogenesis-- 21 days--- /p to the completed
sperm cells
Total 64 days
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perma ocy ogenes s
Among the spermatogonia(all in all*
o'er &illion in &oth testicles) that formthe &asal layer of the germinal epithelium*
se'eral types can &e distinguished
certain type "cells are seen asspermatogonia that di'ide mitotically and
reproduce themsel'es where &y the
spermatogonia population is maintained.
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%he &eginning of spermatogenesis in which the
daughter cells (second group of type " cells)
remain &ound together &y thin bridges ofcytoplasm. %hrough the preser'ation of these
cytoplasmic connections* spermatogonia are
inducted into the spermatogenesis process.
After a further mitotic di'ision type #
spermatogoniaare engendered that also
di'ide themsel'es mitotically into primaryspermatocytes (I).
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%he freshly created primary spermatocytes($)now enter into the first meiosis.
%hey then go immediately into the # phase(that is* into the preleptotenemeiosis)*dou&le their internal DA* lea'e the &asalcompartment and reach the special milieu ofthe luminal compartment.
0ollowing the # phase* these cells attain thecomple+ stage of the prophase of the
meiosisand &ecome there&y noticeablyvisiblewith a light microscope.
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%his prophase* which lasts %4 days* can &e
di'ided into fi'e sections
&eptotene 'ygotene
Pachytene
Diplotene
Diainesis
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In the prophase in e'ery germ cell a
new com&ination of maternal and
paternal genetic material occurs.
After the long prophase follow the
metaphase* anaphase and telophasethat ta3e much less time.
4ne primary spermatocyte yields two
secondary spermatocytes.
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%he secondary spermatocytesgo directly intothe second meiosis* out of which the spermatidsemerge.
#ince in the secondary spermatocytes neitherDA reduplication nor a recom&ination of thegenetic material occurs* the second meiosis canta3e place 5uic3ly.
It lasts only around fi'e hours and for that reasonsecondary spermatocytes are rather seldom seenin a histological section.
%hrough the di'ision of the chromatids of a
secondary spermatocyte* two haploid spermatidsarise that contain only half the original DAcontent.
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esides the sperm cells the
spermatidsare the smallest cells of
the germinal epithelium. In a processlasting se'eral wee3s (so-called
spermiogenesis or
spermiohistogenesis) they are
transformed into spermatocytes with
the acti'e assistance of the #ertoli6scells.
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spermatogenesis wa'e
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As in the diagram* spermatogenesiswa'es mo'e in spirals - li3e a
cor3screw - towards the inner part ofthe lumen.
4utside* on the edge of the tu&uleand at the &eginning of the spiral* liethe spermatogonia7 and* at the end of
the spiral* the fully de'eloped spermcells are in the lumen.
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perm ogenes s(spermatohistogenesis) and
structure of the sperm cell
%he differentiation of the spermatids intosperm cells is called spermiogenesis. It
corresponds to the final part of
spermatogenesis and comprises the
following indi'idual processes that partially
proceed at the same time
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Nuclear condensation thic3ening andreduction of the nuclear si$e*
condensation of the nuclear contents intothe smallest space.
"crosome formation 0orming a cap(acrosome) containing en$ymes that playan important role in the penetrationthrough the pellucid $one of the oocyte.
lagellum formation generation of the
sperm cell tail. *ytoplasma reduction elimination of all
unnecessary cytoplasm.
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.A+onemal structure*
first flagellar
primordium
2.8olgi comple+
9.Acrosomal 'esicle
1.!air of centrioles
(distal and pro+imal):7,itochondrion
.ucleus
;.0lagellar primordium
.Acrosomal cap
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Nuclear condensation%he nucleus &ecomes smaller* denser and
ta3es on a characteristic* flattened form.#een from a&o'e* the nucleus is o'al and*from the narrow side* is pear-shaped.
%he acrosome lies o'er the tip. ucleus andacrosome form the sperm cell6s head that is&ound to the mid-piece &y a short nec3.
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"crosome formation%he 8olgi comple+ engender the
'esicles* which then merge into alarger formation that settles close tothe cell nucleus and finally in'erts itself
li3e a cap o'er the largest part of thenucleus. %he acrosome correspondsfunctionally to a lysosome and thuscontains lysosomal en$ymes(hyaluronidase among others).
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Development of theflagellum%he future a+onemal structuregrows out of one centriole
(distal). %his consists of a&undle of nine peripheraldou&le microtu&ules and two
single ones in the center.
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0our parts of the finished flagellum can &edistinguished
%he neccontains the two centrioles (pro+imal
and distal) among other things. %he mid piececonsists of a sheath of ring-
shaped mitochondria grouped around thea+oneme to pro'ide the energy for the flagellarmo'ement.
%he principle piecehas a sheath of ring fi&ersaround the a+oneme.
%he tailconsists of only the =?2 structure of thea+oneme
%he mature sperm cell is appro+imately > mmlong and completely en'eloped &y the plasmamem&rane.
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. . !lasma mem&rane2.4uter acrosomal mem&rane9.Acrosome1.Inner acrosomal mem&rane
:.ucleus.!ro+imal centriole;.@est of the distal centriole.A+oneme.Anulus2.@ing fi&ers
A.Head.ec3.,id pieceD.!rincipal pieceE.Endpiece%he mature sperm cell is slender7 in the middle part* the mitochondria are thic3 andring-shaped. %he DA in the nucleus is ma+imally condensed.
&e dig+s interstitial cells and
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&eydig+s interstitial cells and
hormonal regulation
etween the seminal canals lie Beydig6sinterstitial cells. %hese are endocrine cells thatmainly produce testosterone* the male se+ualhormone* and release it into the &lood and intothe neigh&oring tissues.
An initial acti'e stage of these cells occurs duringthe em&ryonic de'elopment of the testis. Bater in
Cu'enile life* due to the influence of the &,(luteini$ing hormone) secreted &y the anteriorhypophysis (pituitary gland)* Beydig6s interstitialcells enter a second* long lasting stage of acti'ity.
%ogether with the hormones secreted &y theadrenal corte+* testosterone initiates pu&erty andthus the maturation of the sperm cells.
es os erone pro uc on s rec e y
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es os erone pro uc on s rec e y(luteini$ing hormone)* secreted &y the anterior
lo&e of the hypophysis.
%here &y testosterone can &e transported &y
#ertoli6s cells into the luminal compartment
and there &e concentrated. %estosterone isdecisi'e for spermatogenesis.
%estosterone is also carried away 'ia &loodand lymph fluid. %estosterone has effects on all
tissues* especially also on the &rain during
de'elopment as well as on the se+ual organs.
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4ogenesis
0ollowing the immigration of the primordial germcellsinto the gonadal ridge* they proliferate* are
en'eloped &y coelomic epithelial cells* and form
germinal cords that * though* 3eep their connection
with the coelom epithelium. ow a cortical $one (corte+ o'arii) and a medulla can
&e distinguished* where&y it should &e mentioned that
in females the germinal cords ne'er penetrate into the
medullary $one.
In the genital primordium the following processes then
ta3e place
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A wave of proliferation&egins that lasts
from the :th wee3 to the ;th month
primary germ cells arise in the cortical$one 'ia mitosis of oogonia clones*
&ound together in cellular &ridges* that
happens in rapid succession.
%he cell bridgesare necessary for a
synchronous onset of the su&se5uentmeiosis.
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ith the onset of the meiosis (earliest
onset in the prophase in the 2th wee3)
the designation of the germ cells changes. %hey are now called primary oocytes.
%he primary oocytes &ecome arrested in
the diplotene stage of prophase I (theprophase of the firstmeiotic di'ision).
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#hortly &efore &irth* all the fetal oocytes in thefemale o'ary ha'e attained this stage. %hemeiotic resting phase that then &egins is called
the dictyoteneand it lasts till puberty* duringwhich each month (and in each month thereafteruntil menopause) a pair of primary oocytescomplete the first meiosis.
4nly a few oocytes (secondary oocytes plus onepolar &ody)* though* reach the second meiosisand the su&se5uent o'ulation. %he remainingoocytes that mature each month &ecome atretic.
%he primary oocytes that remain in the o'ariescan stay in the dictyotene stage up tomenopause* in the e+treme case* without e'ermaturing during a menstrual cycle.
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hile the oogonia transform into primary
oocytes* they &ecome restructured so that at
the end of prophase I (the time of thedictyotene) each one gets en'eloped &y a
single layer of flat* follicular epithelial cells
(descendents of the coelomic epithelium).
(oocyte ? follicular epithelium " primordial
follicle).
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4n the one hand* the female germ cell
that at &irth is called the primary oocyte*
and which can de'elop further only during(and after) pu&erty (hormonal cycle is
necessary).
4n the other hand* the follicular epitheliumthat can de'elop further from the
primordial follicle 'ia se'eral follicle
stageswhile oocytes remain in their
primary state.
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%he developmental se-uenceof
the female germ cells is as
follows
Primordial germ cell .
oogonium . primary oocyte .primary oocyte in the
dictyotene
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%he developmental se-uenceof a
follicle goes through 'arious follicle
stages Primordial follicle . primary follicle .
secondary follicle . tertiary follicle
(graafian follicle)
#ince a follicle can die at any moment in
its de'elopment (" atresia)* not all reachthe tertiary follicle stage.
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!tructure of the ovary
An o'ary is su&di'ided into cortical(o'arian corte+) and medullarycompartments(o'arian medulla).
oth &lood and lymph 'essels are found inthe loose connecti'e tissue of the o'arianmedulla.In the cortical compartmentthe oocytes
are present within the various folliclestages.
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%he se+ hormones influence the primordial follicles togrow and a restructuring to ta3e place.
0rom the primordial follicles the primary follicles*
secondary follicles* and tertiary follicles de'elop in turn. 4nly a small percentageof the primordial follicles reach
the tertiary follicle stage - the great maCority meet theirend &eforehand in the 'arious maturation stages.
Barge follicles lea'e scars &ehind in the cortical
compartment and the small ones disappear without atrace.
%he tertiary follicles get to &e the largest and* shortly
&efore o'ulation* can attain a diameter up to 2.: mmthrough a special spurt of growth. %hey are then termedgraafian follicles.
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.!rimordial follicle
2.!rimary follicle
9.#econdary follicle
1.%ertiary follicle:.Antrum folliculi
.umulus oophorus
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%he follicle stages from primordial
follicle to tertiary follicle
Primordial follicle
At the time of birthall the sur'i'ing primary
oocytes are surrounded &y thin/ single layers
of so-called follicular epithelial cells. %hese aredelimited from the rest of the o'arian stroma &y
a thin &asal lamina.
0ollicular epithelial cells are former coelomic
epithelial cells. %he primordial follicles alwaysform the ma0ority of the follicles in the ovary.
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/nder the influence of the se+ hormones
some of themare a&le to develop
furtherto one or more. Although this further de'elopment can
already ta3e place sporadically in the time
&efore &irth and up to pu&erty* the mainpart occurs as soon as a regular hormonal
cycle is esta&lished.
!articularly the last phase of thematuration of a tertiary follicle to &ecome a
large follicle* ready to rupture* remains
reser'ed for the time of regular cycles.
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Primary follicle
In the transition of theprimordial follicles into
primary follicles.
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A.!rimordial follicle
.!rimary follicle
.4ocyte2.0ollicular epithelium
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!econdary folliclehen primary follicles sur'i'e* secondary follicleswith follicular epitheliums encompassing multiple
rowsare engendered. %heir identifying characteristic is a fluid-filled ca'ity*the antral follicle.
%his is now called the stratum granulosum. In thesecondary follicles a glycoprotein layer* the pellucid
$one* &etween the oocyte and follicular epithelium&ecomes 'isi&le.
ytoplasmic processes of the granulosa cells that lieupon it reach the oocyte through the pellucid 1oneand there&y assure their maintenance function.4utside the &asal lamina the stroma o'arii organi$esitself to &ecome theca folliculi cells.
#econdary follicle
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.4ocyte2.!ellucid $one9.#tratum granulosum
1.%heca folliculi cells
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Tertiary follicleIf the secondary follicles sur'i'e* tertiary folliclesare engendered.
%he oocyte lies at the edge in a mound made ofgranulosa epithelial cells* the cumulusoophorus.
In the meantime it has grown so large that itscellular nucleus has attained the si$e of a wholeprimordial follicle.
%he connecti'e tissue around the follicle hasalready clearly differentiated itself into a theca
interna* well supplied with capillaries* out oflarge* lipid-rich cells (hormone production) and atheca e+terna* which forms a transition to thestroma o'arii and contains larger 'essels.
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.4ocyte2.!ellucid $one9.#tratum granulosum
1.%heca interna:.%heca e+terna.Antral follicle;.umulus oophorus (8ranulosa cells*
together with the oocyte)
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Decisi'e for a successful folliclegrowth is a well-de'eloped net of
capillaries in the theca interna. %he precise steering mechanism that
leads to the selection of a follicle and
its su&se5uent maturation to &ecomea graafian follicle is still un3nown.
efore o'ulation a growth spurt of the
tertiary follicles ta3es place.
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2raafian follicle
%his corresponds to anespecially large tertiary folliclethat can &e e+pected to suffice
for o'ulation.
Phase ":
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Phase ":!rimordial germ cells grow* proliferate and &ecome sheathed with coelomicepithelial cells. 8onadal cords arise7 th to
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4ne terms the decline or theregression of follicles of each stage at
e'ery time in the life of a womanfollicular atresia.
%hese follicles do not o'ulate and the
name is deri'ed from that fact. 0ollicle atresia occurs more
intensely* though* at certain moments
(fetal period* early postnatal* &egin ofthe menarche).
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3planations for the onset of heightened atresiaare:
a) in the 54th wee:
Already during the meiotic prophase* mainly in thepachytene stage(lasts the longest* around 9wee3s)* the cells are especially suscepti&le andsuccum&.
ith the formation of the primordial follicle in the
th wee3 the follicular atresiaalso &egins as anadditional reason for the decline in the num&er ofgerm cells.
oth processes together cause the germ cell countto shrin3 to a third (somewhat o'er 2 million o'ary)of the ma+imum num&er.
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postnatal:During the fetal period se+ hormones are producedin the placenta. %his results in a high estrogen le'el
in the &lood of the mother and of the fetus. %his gi'es rise to a considera&le maturation of the
primordial follicles up to the tertiary follicle phase inthe female fetus.
hen the se hormone level in the fetus sinsafter &irth all of the pre'iously matured follicles&ecome atretic (a slight withdrawal &leeding cane'en occur in the newly &orn from the :th to the>th day).
!er o'ary there are then less than 2 million germcells present afterwards.
during puberty:ith th t f & t ( t d 2 f )
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ith the onset of pu&erty (at around 2 years of age) anelevated production of estrogenoccurs again* whichleads to a maturation of the inner and outer genderattri&utes.
After pu&erty is past* around 2:>6>>> germ cells proo'ary remain. ith the onset of a regular cycle a nearlylinear decline commences that with 1> years of ageincreases.
ith the continuous decrease of the follicle cell count theproduction of estrogen is also reduced constantly. ithroughly :> years of age when follicles are no longerpresent* the estrogen production ceases andmenopause ensues
It is assumed the dictyotene stage of meiosis representsa special condition for oocytes that e+hi&its great sta&ilitywith regard to outer physical and chemical influences. Incontrast* the earlier stages* especially the pachytene*are more sensiti'e.
Th i l
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The ovarian cycle
4f the roughly :>>6>>> follicles that are
present in the two o'aries at the &eginning
of se+ual maturity* only around 1 reach
the graafian follicle stage and are thusa&le to release oocytes (o'ulation). %his
num&er is simply deri'ed &y multiplying
the num&er of cycles per year (2) and thenum&er of years in which a woman is
fertile (1>).
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I th t f thi h l t l i th
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In the centerof this hormonal control is thehypothalamamics.hypophysial (pituitarygland) systemwith the two hypophysial
gonadotropins 0#H and BH. %he pulsating li&eration of 8n@H &y the
hypothalamus is the fundamental preconditionfor a normal control of the cyclic o'arian
function. %his cyclic acti'ity releases 0#H and BH* &oth of
which stimulate the maturation of the follicles inthe o'ary and trigger ovulation.
During the o'arian cycle* estrogen is produced&y the theca interna and follicular cells (in theso-called follicle phase) and progesterone &y thecorpus luteum (so-called luteal phase).
%h t l i it f th h l l
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%he control circuit of the hormonal cycle
has two essential control elements
%he pulsatileli&eration of 8n@H* as wellas 0#H and BH
As a rule the ovarian cycle lasts %
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As a rule* the ovarian cycle lasts %
days. It is su&di'ided into two phases
ollicle phase recruitmentof a so-calledfollicle cohort and* within this* the selection
of the mature follicle. %his phase ends
with ovulation.
3stradiolis the steering hormone.
ormally* it lasts 1 days* &ut this can
vary considerably7
&uteal phase progesteron production
&y the yellow &ody (" corpus luteum)
and lasts 1 days (relati'ely constant).
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A.0ollicle phase.Buteal phase
.!rimary follicle
D.#econdary follicleE.%ertiary follicle0.8raafian follicle
E2 Estradiol
!r !rogesteroneBH Buteini$ing hormone0#H 0ollicle stimulating hormone
ll % ll Di i i ! d t
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ell %ype ell Di'ision !roduct
#permatagonium (2)/ndergoes ,itosis
!rimary #permatocyte (2) !rimary #permatocyte (2)/ndergoes
,eiosis I F results in
2 #econdary #permatocytes () #econdary #permatocyte ()/ndergoes
,eiosis II* results in #permatid ().
#permatid ()--,aturation - no cell di'ision--#perm cell ()
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4ogonium (2)de'elopment - no cell di'ision--!rimary 4ocyte (2).
!rimary 4ocyte (2)/ndergoes ,eiosis I ---#econdary 4ocyte () and 0irst !olar ody().
#econdary 4ocyte ()/ndergoes ,eiosis II(stimulated &y fertili$ation from sperm)-----4'um () and #econd !olar ody ()