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Reproduction in Domestic Animals

Reproduction in Domestic Animals

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Reprod Dom Anim 43 (Suppl. 2), 245–251 (2008); doi: 10.1111/j.1439-0531.2008.01169.xISSN 0936-6768Fertilization and Early Embryonic Development <strong>in</strong> the Porc<strong>in</strong>e Fallopian TubeK-P Bru¨ssow 1 ,JRátky 2 and H Rodriguez-Mart<strong>in</strong>ez 31 Department of Reproductive Biology, FBN Research Institute for the Biology of Farm <strong>Animals</strong>, Dummerstorf, Germany; 2 Department of Biology of<strong>Reproduction</strong>, Research Institute for Animal Breed<strong>in</strong>g and Nutrition, Herceghalom, Hungary; 3 Division of <strong>Reproduction</strong>, Faculty of Veter<strong>in</strong>aryMedic<strong>in</strong>e and Animal Science, Swedish University of Agricultural Sciences (SLU), Uppsala, SwedenContentsFertilization and early embryo development relies on acomplex <strong>in</strong>terplay between the Fallopian tube and thegametes before and after fertilization. Thereby the oviduct,as a dynamic reproductive organ, enables reception, transportand maturation of male and female gametes, theirfusion, and supports early embryo development. This paperreviews current knowledge regard<strong>in</strong>g physiological processesbeh<strong>in</strong>d the transport of boar spermatozoa, their storage <strong>in</strong>and release from the functional sperm reservoir (SR), and ofthe <strong>in</strong>teractions that newly ovulated oocytes play with<strong>in</strong> thetube dur<strong>in</strong>g their transport to the site of fertilization.Experimental evidence of an ovarian control on spermrelease from the SR is highlighted. Furthermore, the impactof oviductal secretion on sperm capacitation, oocyte maturation,fertilization and early embryo development isstressed.IntroductionThe oviduct (synonyms Fallopian tube, salp<strong>in</strong>x, uter<strong>in</strong>etube) firstly described by Gabriel Fallopius <strong>in</strong> 1561, wasconsidered a simple connection between the ovary andthe uterus for a long time. Today, the oviduct isregarded as one of the most dynamic reproductiveorgans, whose functional aspects are not yet fullyunderstood (Rodriguez-Mart<strong>in</strong>ez et al. 2001). The oviductprovides a favourable environment for spermatozoa,enabl<strong>in</strong>g their transport and ensur<strong>in</strong>g beneficialconditions dur<strong>in</strong>g storage while prepar<strong>in</strong>g them forfertilization; as well as oocytes, mak<strong>in</strong>g ovum pick-upand transport to the site of fertilization possible, whileensur<strong>in</strong>g their f<strong>in</strong>al maturation. Both fertilization andearly embryonic development take place with<strong>in</strong> theoviduct. This paper reviews physiological processesbeh<strong>in</strong>d the transport of boar spermatozoa, their storage<strong>in</strong> and release from the functional sperm reservoir (SR),and of the <strong>in</strong>teractions that newly ovulated oocytes playdur<strong>in</strong>g their transport to the site of fertilization. Someaspects are highlighted, such as experimental evidencefor an ovarian <strong>in</strong>fluence on sperm release from the SRand the relevance of oviductal secretion on spermcapacitation, oocyte maturation, fertilization and earlyembryo development.The OviductThe porc<strong>in</strong>e oviduct is approximately 25 cm <strong>in</strong> length(Bru¨ ssow 1985) and it is anatomically divided <strong>in</strong>to threema<strong>in</strong> segments, e.g. the <strong>in</strong>fundibulum, the ampulla andthe isthmus. Connect<strong>in</strong>g areas, i.e. the term<strong>in</strong>al section(ostium), the ampulla-isthmic-junction (AIJ) and theutero-tubal-junction (UTJ) are also dist<strong>in</strong>guished. Thehistoarchitecture of the Fallopian tube is simple, witha non-glandular mucosa (endosalp<strong>in</strong>x), covered with al<strong>in</strong><strong>in</strong>g epithelium with secretory and ciliated cells,a double-layered smooth muscle (myosalp<strong>in</strong>x) and acover<strong>in</strong>g serosa (mesosalp<strong>in</strong>x). Towards the ostium, thethickness of the <strong>in</strong>ternal, circular muscle becomesth<strong>in</strong>ner; the longitud<strong>in</strong>al mucosal plicae ga<strong>in</strong> complexityand the number of ciliated cells considerably<strong>in</strong>creases (Rodriguez-Mart<strong>in</strong>ez et al. 2001; Yaniz et al.2006). This histoarchitecture builds tubal compartmentswith apparently different roles, all with optimal environmentwith regards to pH, osmotic pressure, nutrients,specific secretory products and signal molecules(reviewed by Rodriguez-Mart<strong>in</strong>ez 2007). Thus, preparationof gametes, fertilization and early embryonicdevelopment are supported and regulated by theoviduct.Sperm Transport and the Tubal SpermReservoirAfter deposition of spermatozoa <strong>in</strong> the cervix by mat<strong>in</strong>gor AI, a sperm subpopulation is rapidly transportedthrough the uterus, whereas the majority is elim<strong>in</strong>atedfrom the uter<strong>in</strong>e lumen (Rodriguez-Mart<strong>in</strong>ez et al.2005). Spermatozoa that ascended the uterus <strong>in</strong> the firstphase of sperm transport colonize the UTJ and thecaudal isthmus (>10 4 spermatozoa), a segment thatacts as a functional pre-ovulatory sperm reservoir,temporarily arrest<strong>in</strong>g spermatozoa (up to 30 h) and,presumably, activat<strong>in</strong>g them at a given time (Rodriguez-Mart<strong>in</strong>ez et al. 2005). In the SR, most spermatozoama<strong>in</strong>ta<strong>in</strong> normal ultrastructure and viability (Rodriguez-Mart<strong>in</strong>ezet al. 1990; Mburu et al. 1997). Severalconcerted factors are thought to expla<strong>in</strong> the formationof the functional SR, <strong>in</strong>clud<strong>in</strong>g the narrowed lumen(Hunter 1984), viscous mucus (Johansson et al. 2000),lower temperature (Hunter and Nichol 1986), localenzymatic and ionic milieu (Rodriguez-Mart<strong>in</strong>ez et al.1991), selective b<strong>in</strong>d<strong>in</strong>g of spermatozoa to the epithelium(Fazeli et al. 1999), and specific tubal fluid components(Tienthai et al. 2004), all which primarily leadto sperm quiescence.The sequential release of a restricted number ofspermatozoa from the SR towards the AIJ ensuresfertilization of oocytes with<strong>in</strong> a time w<strong>in</strong>dow, even ifovulation lasts over a longer <strong>in</strong>terval. The periovulatoryprogression of spermatozoa is suggested to be a complexand concerted process <strong>in</strong>clud<strong>in</strong>g open<strong>in</strong>g of the lumenby a decrease <strong>in</strong> the hormonally-driven endosalp<strong>in</strong>gealoedema, dissolution of the hyaluronan (HA)-richÓ 2008 The Authors. Journal compilation Ó 2008 Blackwell Verlag

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