sperm oogenesis

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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 ()