Male Reproductive System - Germ Line

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As in the hermaphrodite, the male germ line exhibits distal-proximal polarity with a mitotic cell population located at the most distal end of the gonad (DG) and meiotic cells extending proximally. Gametogenesis occurs in the proximal germ line and the resulting spermatids are stored in the seminal vesicle until ejaculation (<a href="#MaleReproFIG17A">MaleReproFIG 17A</a>;<a href="#Hirsh1976"> Hirsh et al., 1976</a>; <a href="#Klass1976">Klass et al., 1976</a>; <a href="#Kimble1979">Kimble and Hirsh, 1979</a>). </p> <div align="center"><img src="images/MaleReproFIG17A.jpg" alt="MaleReproFIG 17A Adult male germ line organization" name="MaleReproFIG17A" width="750" height="278" border="0" align="middle" class="Galborder" id="MaleReproFIG17A"><a name="MaleReproFIG17A" id="MaleReproFIG17A"></a></div> <span class="picturetitle"> MaleReproFIG 17A: Adult male germ line organization. </span><span class="style2">Nomarski DIC picture of an adult male tail region featuring the parts of the reproductive tract, lateral view. (DG) Distal gonad; (PG) proximal gonad; (DTC) distal tip cell.</span> <p>Much of the male germ line is a syncytium. Germ cells have incomplete borders and are connected to one another via a central canal, the rachis (<a href="#MaleReproFIG17B">MaleReproFIG 17B</a> and <a href="#MaleReproFIG17C">17C</a>). The germ line is ensheathed only by the gonadal basal lamina (gbl) which also covers the somatic portion of the gonad. </p> <div align="center"><img src="images/MaleReproFIG17B.jpg" alt="MaleReproFIG 17B Ultrastructure of the male germ line" name="MaleReproFIG17B" width="480" height="586" border="0" align="middle" class="Galborder" id="MaleReproFIG17B"><a name="MaleReproFIG17B" id="MaleReproFIG17B"></a></div> <span class="picturetitle">MaleReproFIG 17B: Ultrastructure of the male germ line. </span><span class="style2">TEM shows cross section of both distal and proximal arms of the germ line, transverse view. (Image source: him5-25 [Hall] 9112.) (Gbl) Gonadal basal lamina. </span> <div align="center"><br> <img src="images/MaleReproFIG17C.jpg" alt="MaleReproFIG 17C Ultrastructure of the male germ line" name="MaleReproFIG17C" width="478" height="526" border="0" align="middle" class="Galborder" id="MaleReproFIG17C"><a name="MaleReproFIG17C" id="MaleReproFIG17C"></a></div> <span class="picturetitle style2"> MaleReproFIG 17C: </span><span class="picturetitle">Ultrastructure of the male germ line. </span><span class="style2">TEM shows cross section of proximal arm of the germ line lying adjacent to the intestine, transverse view. (Image source: him5-25 [Hall] 0702.) (Gbl) Gonadal basal lamina. </span> <hr class="page-splits"> <p><span class="subtitle"><a name="Mitosisiinthegermline2" id="Mitosisiinthegermline2"></a></span><span class="style10"><br> 2 Mitosis in the Germ Line </span></p> <p>As in the hermaphrodite, the distal end of the male germ line (<a href="#MaleReproFIG18">MaleReproFIG 18A</a>) contains a stem cell population and is referred to as the mitotic zone. As germ cells move away from the influence of the distal tip cell (DTC, described in <a href="../somaticgonad/Somaticgonadframeset.html" target="_blank">Male Reproductive System - Somatic Gonad</a>), they enter meiosis I, proceeding through prophase I to pachytene (<a href="#Hirsh1976">Hirsh et al., 1976</a>). Nuclear morphology at various stages of mitosis and meiosis I can visualized with DAPI staining (examples from the hermaphrodite germ line, highlighting stage-specific characteristics, are shown in <a href="#MaleReproFIG18">MaleReproFIG 18B-D</a>).</p> <div align="center"><img src="images/MaleReproFIG18.jpg" alt="MaleReproFIG 18 Cell division in the germ line" name="MaleReproFIG18" width="707" height="546" border="0" align="middle" class="Galborder" id="MaleReproFIG18"><a name="MaleReproFIG18" id="MaleReproFIG18"></a> </div> <span class="picturetitle"> MaleReproFIG 18: Cell division in the germ line.</span><span class="style2"><strong> A. </strong>Nomarski DIC image shows locations of mitosis, meiosis and gametogenesis in the germ line. <strong>B-D. </strong>DAPI -stained germ line cells. (Image source: J. Maciejowski and E.J. Hubbard). <strong>B. </strong>Mitosis occurs at the distal end of the germ line in an area called the mitotic zone. <strong>C.</strong> As germ cells move farther away from the distal tip cell (DTC), they enter an area called the transition zone where they enter meiosis I. <strong>D. </strong>Cells proceed through prophase I to pachytene. Chromosomes spread out along the periphery of the nucleus forming a "bowl of spaghetti" phenotype. </span> <hr class="page-splits"> <p><span class="subtitle"><a name="Spermatogenesisandspermiogenesis3" id="Spermatogenesisandspermiogenesis3"></a></span><span class="style10"><br> 3 Spermatogenesis and Spermiogenesis</span> </p> <p>After pachytene, the precursor of spermatids, the spermatocytes, detach from the rachis and complete meiosis generating haploid spermatids (<a href="#MaleReproFIG19">MaleReproFIG 19</a> and <a href="#MaleReproFIG20">20</a>). This process of spermatid formation is called spermatogenesis. Spermatids are stored in the seminal vesicle until ejaculation. Spermiogenesis, activation of sessile spermatids into crawling spermatozoa, occurs after mating, in the hermaphrodite uterus. Some spermatids that remain in the male after mating, however, may mature into spermatozoa and can be seen swimming around the vas deferens and seminal vesicle (<a href="#Ward1979">Ward and Carrel, 1979</a>). Recent studies suggest that the male gonad contains substances that prevent premature activation of spermatids prior to sperm transfer (<a href="#Stanfield2006">Stanfield and Villeneuve, 2004</a>).</p> <div align="center"><img src="images/MaleReproFIG19.jpg" alt="MaleReproFIG 19 Sperm development" name="MaleReproFIG19" width="514" height="313" border="0" align="middle" class="Galborder" id="MaleReproFIG19"><a name="MaleReproFIG19" id="MaleReproFIG19"></a> </div> <span class="picturetitle"> MaleReproFIG 19: Sperm development. </span><span class="style2">Illustration showing the stages of spermatogenesis and spermiogenesis resulting the production of motile sperm. (Based on <a href="#LHernault1997">L'Hernault, 1997</a>.) </span> <hr class="page-splits"></td> </tr> <tr> <td align="left" background="#Chinsang2000" class="style1"> <div align="center"> <img src="images/MaleReproFIG20.jpg" alt="MaleReproFIG 20 Male sperm production" name="MaleReproFIG20" width="409" height="323" border="0" align="middle" class="Galborder" id="MaleReproFIG20"><a name="MaleReproFIG20" id="MaleReproFIG20"></a> </div> <span class="picturetitle"> MaleReproFIG 20: Male sperm production. </span><span class="style2">Nomarski DIC picture showing the location of spermatocytes and spermatids within the adult male germ line, lateral view. </span> <p>Developing spermatocytes contain a large number of specialized vesicles called FB-MOs (<em>F</em>ibrous <em>B</em>ody-<em>M</em>embranous <em>O</em>rganelle) (<a href="#MaleReproFIG21">MaleReproFIG 21</a>, <a href="#MaleReproFIG22">22</a> and <a href="#MaleReproFIG24">23</a>). These organelles contain proteins required in the future spermatid and spermatozoon, including MSP (<em>M</em>ajor <em>S</em>perm <em>P</em>rotein; <a href="#Wolf1978">Wolf et al., 1978</a>; <a href="#Ward1982">Ward and Klass,1982</a>). During development, the FB-MOs partition with the portion of the spermatocyte destined to become the future spermatid (<a href="#Ward1986">Ward, 1986</a>). The residual body (<a href="#MaleReproFIG22">MaleReproFIG 22</a>) acts as a depot for proteins and organelles no longer required by the developing spermatid (<a href="#LHernault1997">L'Hernault, 1997</a>; <a href="#Arduengo1998">Arduengo et al., 1998</a>; <a href="#Kelleher2000">Kelleher et al., 2000</a>). </p> <div align="center"><img src="images/MaleReproFIG21.jpg" alt="MaleReproFIG 21 FB-MOs production" name="MaleReproFIG21" width="750" height="225" border="0" align="middle" class="Galborder" id="MaleReproFIG21"><a name="MaleReproFIG21" id="MaleReproFIG21"></a> </div> <span class="picturetitle"> MaleReproFIG 21: FB-MOs production. </span><span class="style2">Panel on left shows the process of FB-MO (fibrous body membranous organelle) formation. Right panel illustrates the parts of the FB-MO. (Based on <a href="#Arduengo1998">Arduengo et al., 1998</a>.) </span> <p>Maturing spermatids and spermatozoa have highly condensed nuclei (N) and tightly packed mitochondria (M) (<a href="#MaleReproFIG23">MaleReproFIG 23</a>). In spermatids, MOs (now lacking FB) locate near the cell periphery. During spermatid activation, MOs fuse with the plasma membrane releasing their contents (primarily glycoproteins) onto the cell surface. A fusion pore is generated on the cell surface by the MO collar (<a href="#MaleReproFIG24">MaleReproFIG 24</a>). Mutants affected in MO fusion produce sperm with defective motility suggesting that MO content enhances sperm mobility (<a href="#Ward1981">Ward et al., 1981</a>; <a href="#Roberts1986">Roberts et al, 1986</a>;<a href="#Achanzar1997"> Achanzar and Ward, 1997</a>). Spermatid activation also involves the formation of a foot or pseudopodium (<a href="#MaleReproFIG24">MaleReproFIG 24</a>). In contrast to sperm in other phyla, <em>C. elegans</em> sperm lack flagella. Pseudopodia allow spermatozoa to attach to the walls of the spermathecal or seminal vesicle lumen and to crawl. This motility is driven by dynamic polymerization of MSP (Major Sperm Protein). In addition to an intracellular cytoskeletal function, MSP contains sequences that mediate extracellular signaling (see Oocyte maturation section of <a href="../../hermaphrodite/germ line/Germframeset.html" target="_blank">Hermaphrodite Reproductive System - Germ Line</a>; <a href="#Miller2001">Miller et al, 2001</a>; <a href="#Italiano1996">Italiano et al., 1996</a>; <a href="#Roberts2000">Roberts and Stewart, 2000</a>). <span class="style2"><br> </span> </p> <div align="center"> <img src="images/MaleReproFIG22.jpg" alt="MaleReproFIG 22 Ultrastructure of FB-MO" name="MaleReproFIG22" width="424" height="328" border="0" align="middle" class="Galborder" id="MaleReproFIG22"><a name="MaleReproFIG22" id="MaleReproFIG22"></a> </div> <span class="picturetitle"> MaleReproFIG 22: Ultrastructure of FB-MO. </span><span class="style2">TEM of FB-MO in developing spermatocytes, transverse view. (Image source: him5-25 [Hall] 1215.) (SV) Seminal vesicle. </span> <hr class="page-splits"> <div align="center"><img src="images/MaleReproFIG23.jpg" alt="MaleReproFIG 23 Ultrastructure of FB-MO in maturing sperm" name="MaleReproFIG23" width="529" height="328" border="0" align="middle" usemap="#MaleIntroFIG6Map" class="Galborder" id="MaleReproFIG23"><a name="MaleReproFIG23" id="MaleReproFIG23"></a> <span class="picturename"> </span></div> <span class="picturetitle"> MaleReproFIG 23: Ultrastructure of FB-MO in maturing sperm. </span><span class="style2">TEM of FB-MO in maturing spermatids and spermatozoa showing the highly condensed nuclei and tightly packed mitochondria, transverse view. The collar region of the FB-MO is indicated by <em>white arrows</em>. (Image source: him5-25 [Hall] 2631.) (SV) Seminal vesicle. </span> <hr class="page-splits"></td> <tr> <td align="left" background="#Chinsang2000" class="style1"><div align="center"><img src="images/MaleReproFIG24.jpg" alt="MaleReproFIG 24 Ultrastructure of spermatozoon" name="MaleReproFIG24" width="450" height="442" border="0" align="middle" class="Galborder" id="MaleReproFIG24"> <a name="MaleReproFIG24" id="MaleReproFIG24"></a></div> <span class="picturetitle">MaleReproFIG 24: Ultrastructure of spermatozoon. </span><span class="style2">TEM of spermatid activation. The MO fuses with the plasma membrane via the collar. Pseudopodia can all be seen. (Image source: him5-25 [Hall] H707.)</span> <hr class="page-splits"> <span class="subtitle"><a name="Germlinedevelopment4" id="Germlinedevelopment4"></a></span><span class="style10"><br> 4 Germ-line Development </span> <p>Male germ line development spans L1 to early adulthood. All germ cells are descended from either Z2 or Z3 (<a href="#Kimble1979">Kimble and Hirsh, 1979</a>). Key events in germ line development are summarized in <a href="#MaleReproFIG25">MaleReproFIG 25</a>. In contrast to somatic lineage development in <em>C. elegans</em>, germ line cell divisions appear to be variable with respect to their timing and planes of division (<a href="#Kimble1979">Kimble and Hirsh, 1979</a><a href="http://elegans.swmed.edu/wli/%5Bcgc193%5D/?%5Cbh%5Bi%ED%EC%EE%EF%5Dr%5Bs%DF%5Dh%5Cb%3B%5Cbk%5Bi%ED%EC%EE%EF%5Dmbl%5Be%E9%E8%EA%EB%5D%5Cb" target="_blank"></a>) and hence the precise lineal relationships between these cells are not known. </p> <div align="center"><img src="images/MaleReproFIG25.jpg" alt="MaleReproFIG 25 Male germ line development" name="MaleReproFIG25" width="610" height="586" border="0" align="middle" class="Galborder" id="MaleReproFIG25"><a name="MaleReproFIG25" id="MaleReproFIG25"></a></div> <span class="picturetitle">MaleReproFIG 25: Male germ line development. </span><span class="style2">Illustration showing the stages of germ line formation in the developing male. (Gbl) Gonadal basal lamina; (DTC) distal tip cell; (SPm) somatic gonadal primordium of the male; (DG) distal gonad; (PG) proximal gonad. (Adapted from <a href="#Schedl1997">Schedl, 1997</a>.) </span> <hr class="page-splits"> <span class="subtitle"><a name="References5" id="References5"></a></span><span class="style10"><br> 5 References</span> <p><a name="Achanzar1997"></a>Achanzar, W.E. and Ward, S. 1997. A nematode gene required for sperm vesicle fusion. J. Cell Sci. <strong>110: </strong>1073-1081. <a href="http://jcs.biologists.org/content/110/9/1073.long" target="_blank">Article</a></p> <p><a name="Arduengo1998"></a>Arduengo, P.M., Appleberry, O.K., Chuang, P. and L'Hernault, S.W. 1998. The presenilin protein family member SPE-4 localizes to an ER/Golgi derived organelle and is required for proper cytoplasmic partitioning during <em>Caenorhabditis elegans</em> spermatogenesis. J. Cell Sci. <strong>111:</strong> 3645-3654. <a href="http://jcs.biologists.org/content/111/24/3645.long" target="_blank">Article</a></p> <p><a name="Hirsh1976"></a>Hirsh, D., Oppenheim, D. and Klass, M. 1976. Development of the reproductive system of <em>Caenorhabditis elegans</em>. Dev. Biol. <strong>49: </strong>200-219. <a href="http://dx.doi.org/10.1016/0012-1606(76)90267-0" target="_blank">Abstract</a></p> <p><a name="Italiano1996"></a>Italiano, J.E. Jr., Roberts, T.M., Stewart, M. and Fontana, C.A.  1996.  Reconstitution in vitro of the motile apparatus from the amoeboid sperm of <em>Ascaris</em> shows that filament assembly and bundling move membranes. Cell <strong>84:</strong> 105-114. <a href="http://dx.doi.org/10.1016/S0092-8674(00)80997-6" target="_blank">Article</a></p> <p><a name="Kelleher2000"></a>Kelleher, J.F., Mandell, M.A., Moulder, G., Hill, K.L., L'Hernault, S.W., Barstead, R. and Titus, M.A. 2000. Myosin VI is required for asymmetric segregation of cellular components during <em>C. elegans</em> spermatogenesis. Curr. Biol.<strong> 10:</strong> 1489-1496. <a href="http://dx.doi.org/10.1016/S0960-9822(00)00828-9" target="_blank">Article</a></p> <p><a name="Kimble1979"></a>Kimble, J. and Hirsh, D. 1979. The postembryonic cell lineages of the hermaphrodite and male gonads in <em>Caenorhabditis elegans</em>. Dev. Biol. <strong>70</strong>: 396-417. <a href="../../ver1/Postemblingonad_1979/toc.html" target="_blank">Article </a></p> <p><a name="Klass1976"></a>Klass, M., Wolf, N. and Hirsh, D. 1976. Development of the male reproductive system and sexual transformation in the nematode <em>C. elegans.</em> Dev. Biol. <strong>52:</strong> 1-18. <a href="http://dx.doi.org/10.1016/0012-1606(76)90002-6" target="_blank">Abstract</a></p> <p><a name="LHernault1997"></a>L'Hernault, S.W. 1997. Spermatogenesis. In <em>C. elegans II</em> (ed. D. L. Riddle et al.). Chap. 11. pp. 417-500. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. <a href="http://www.ncbi.nlm.nih.gov/books/NBK20101/" target="_blank">Article</a></p> <p><a name="Miller2001"></a>Miller, M.A., Nguyen, V.Q., Lee, M.H., Kosinski, M., Schedl, T., Caprioli, R.M. and Greenstein, D. 2001. A sperm cytoskeletal protein that signals oocyte meiotic maturation and ovulation. Science <strong>291:</strong> 2144-2147. <a href="http://dx.doi.org/10.1126/science.1057586" target="_blank">Abstract</a></p> <p><a name="Roberts2000"></a>Roberts, T.M. and Stewart, M. 2000. Acting like actin. The dynamics of the nematode major sperm protein (msp) cytoskeleton indicate a push-pull mechanism for amoeboid cell motility. J. Cell Biol. <strong>149:</strong> 7-12. <a href="http://dx.doi.org/10.1083/jcb.149.1.7" target="_blank">Article</a></p> <p><a name="Roberts1986"></a>Roberts, T.M., Pavalko, F.M. and Ward, S. 1986. Membrane and cytoplasmic proteins are transported in the same organelle complex during nematode spermatogenesis. J. Cell Biol. <strong>102:</strong> 1787-1796. <a href="http://jcb.rupress.org/content/102/5/1787.long" target="_blank">Article</a></p> <p><a name="Schedl1997" id="Schedl1997"></a>Schedl, T. 1997. Developmental Genetics of the Germ Line. In <em>C. elegans II</em> (ed. D. L. Riddle et al.). chap. 10. pp. 417-500. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York.<a href="http://www.ncbi.nlm.nih.gov/books/NBK20133/" target="_blank"> Article</a></p> <p><a name="Stanfield2006"></a>Stanfield, G.M. and Villeneuve, A.M. 2006. Regulation of sperm activation by SWM-1 is required for reproductive success of <em>C. elegans</em> males. Curr. Biol. <strong>16: </strong>252-63. <a href="http://dx.doi.org/10.1016/j.cub.2005.12.041" target="_blank">Article</a></p> <p><a name="Ward1986"></a>Ward, S. 1986. Asymmetric localization of gene products during the development of <em>C. elegans</em> spermatozoa. In Gametogenesis and the Early Embryo. (ed. J. G. Gall). pp. 55-75. 44th Symposium of the Society for Developmental Biology. Alan R. Liss, New York.</p> <p><a name="Ward1979"></a>Ward, S. and Carrel, J.S. 1979. Fertilization and sperm competition in the nematode <em>Caenorhabditis elegans. </em>Dev. Biol. <strong>73:</strong> 304-321. <a href="http://dx.doi.org/10.1016/0012-1606(79)90069-1" target="_blank">Abstract</a></p> <p><a name="Ward1982"></a>Ward, S. and Klass, M. 1982. The location of the major protein in <em>Caenorhabditis elegans</em> sperm and spermatocytes. Dev. Biol. <strong>92:</strong> 203-208. <a href="http://dx.doi.org/10.1016/0012-1606(82)90164-6" target="_blank">Abstract</a></p> <p><a name="Ward1981"></a>Ward, S., Argon, Y. and Nelson, G.A. 1981. Sperm morphogenesis in wild-type and fertilization-defective mutants of <em>Caenorhabditis elegans.</em> J. Cell Biol. <strong>91:</strong> 26-44. <a href="http://jcb.rupress.org/content/91/1/26.long" target="_blank">Article</a></p> <p><a name="Wolf1978"></a>Wolf, N., Hirsh, D. and McIntosh, J.R. 1978. Spermatogenesis in males of the free-living nematode, <em>Caenorhabditis elegans</em>. J. Ultrastruct. Res. <strong>63:</strong> 155-69. <a href="http://dx.doi.org/10.1016/S0022-5320(78)80071-9" target="_blank">Abstract</a></p> <hr class="page-splits"></td> </tr><tr> <td align="left" class="style1"> <a href="#top"><img src="../../picts/topoff.gif" border="0" onMouseOver="this.src='../../picts/topon.gif'" onMouseOut="this.src='../../picts/topoff.gif'"> </a><br> This chapter should be cited as: Lints, R. and Hall, D.H. 2009. Male reproductive system, germ line. In<em> WormAtlas</em>. doi:<a href="http://10.3908/wormatlas.2.16" target="_top">10.3908/wormatlas.2.16</a><br> Edited for the web by Laura A. Herndon. 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