# Sperm bauplan and function and underlying processes of sperm formation and selection **Authors:** Maria Eugenia Teves, Eduardo R. S. Roldan PMC · DOI: 10.1152/physrev.00009.2020 · 2021-04-21 ## TL;DR This paper explores the structure and function of sperm cells, focusing on the evolutionary and molecular mechanisms behind their diversity. ## Contribution The paper provides a comprehensive analysis of the adaptive and mechanistic origins of sperm diversity. ## Key findings - Sperm cells show significant structural and functional diversity across species. - Selective forces and molecular mechanisms in the testis drive sperm differentiation and variation. - Understanding these mechanisms can explain male reproductive success and evolutionary adaptations. ## Abstract The spermatozoon is a highly differentiated and polarized cell, with two main structures: the head, containing a haploid nucleus and the acrosomal exocytotic granule, and the flagellum, which generates energy and propels the cell; both structures are connected by the neck. The sperm’s main aim is to participate in fertilization, thus activating development. Despite this common bauplan and function, there is an enormous diversity in structure and performance of sperm cells. For example, mammalian spermatozoa may exhibit several head patterns and overall sperm lengths ranging from ∼30 to 350 µm. Mechanisms of transport in the female tract, preparation for fertilization, and recognition of and interaction with the oocyte also show considerable variation. There has been much interest in understanding the origin of this diversity, both in evolutionary terms and in relation to mechanisms underlying sperm differentiation in the testis. Here, relationships between sperm bauplan and function are examined at two levels: first, by analyzing the selective forces that drive changes in sperm structure and physiology to understand the adaptive values of this variation and impact on male reproductive success and second, by examining cellular and molecular mechanisms of sperm formation in the testis that may explain how differentiation can give rise to such a wide array of sperm forms and functions. ## Full-text entities - **Genes:** Spata16 (spermatogenesis associated 16) [NCBI Gene 70862] {aka 4921511F01Rik, 4930503K02Rik, Nyd-sp12}, Spag6 (sperm associated antigen 6) [NCBI Gene 381350] {aka Gm13334, Spag6l}, Ift20 (intraflagellar transport 20) [NCBI Gene 55978] {aka 0610009H04Rik, mIFT20}, Snapin (SNAP-associated protein) [NCBI Gene 20615] {aka 25kDa, Bloc1s7, Snap25bp, Snapap}, DNAH6 (dynein axonemal heavy chain 6) [NCBI Gene 1768] {aka DNHL1, Dnahc6, HL-2, HL2}, Ift88 (intraflagellar transport 88) [NCBI Gene 21821] {aka Tg737, Tg737Rpw, TgN737Rpw, Ttc10, flexo, fxo}, Actr3 (ARP3 actin-related protein 3) [NCBI Gene 74117] {aka 1200003A09Rik, Arp3}, Hspg2 (perlecan (heparan sulfate proteoglycan 2)) [NCBI Gene 15530] {aka HSPG, Pcn, Plc, per}, Myo5a (myosin VA) [NCBI Gene 17918] {aka 9630007J19Rik, Dbv, MVa, Myo5, MyoVA, Sev-1}, PRM2 (protamine 2) [NCBI Gene 5620] {aka CT94.2}, KCNQ1OT1 (KCNQ1 opposite strand/antisense transcript 1) [NCBI Gene 10984] {aka KCNQ1-AS2, KCNQ10T1, Kncq1, KvDMR1, KvLQT1-AS, LIT1}, Pfn3 (profilin 3) [NCBI Gene 75477] {aka 1700012P12Rik}, CATSPER1 (cation channel sperm associated 1) [NCBI Gene 117144] {aka CATSPER, SPGF7}, Syne3 (spectrin repeat containing, nuclear envelope family member 3) [NCBI Gene 212073] {aka KASH3, nesprin-3}, Spef2 (sperm flagellar 2) [NCBI Gene 320277] {aka C230086A09Rik}, Kifc1 (kinesin family member C1) [NCBI Gene 100502766] {aka 100042970, Gm4137, HSET, KNSL2, Kifc5a, Knsl2a}, SUN3 (Sad1 and UNC84 domain containing 3) [NCBI Gene 256979] {aka SUNC1}, Rimbp3 (RIMS binding protein 3) [NCBI Gene 239731] {aka Gm1759, Gm603, RIM-BP3, mKIAA1666}, Syne1 (spectrin repeat containing, nuclear envelope 1) [NCBI Gene 64009] {aka 8B, A330049M09Rik, C130039F11Rik, CPG2, KASH1, Myne1}, Tnp2 (transition protein 2) [NCBI Gene 21959] {aka TP2, Tp-2}, Spag17 (sperm associated antigen 17) [NCBI Gene 74362] {aka 4931427F14Rik, PF6, Spag17-ps}, CA2 (carbonic anhydrase 2) [NCBI Gene 760] {aka CA-II, CAC, CAII, Car2, HEL-76, HEL-S-282}, SYNE1 (spectrin repeat containing nuclear envelope protein 1) [NCBI Gene 23345] {aka 8B, AMC3, AMCM, ARCA1, C6orf98, CPG2}, Rnf19a (ring finger protein 19A, RBR E3 ubiquitin protein ligase) [NCBI Gene 362900] {aka Rnf19}, Lbr (lamin B receptor) [NCBI Gene 98386] {aka C14SR, ic}, CAMP (cathelicidin antimicrobial peptide) [NCBI Gene 820] {aka CAP-18, CAP18, CRAMP, FALL-39, FALL39, HSD26}, Ranbp17 (RAN binding protein 17) [NCBI Gene 66011], Pafah1b1 (platelet-activating factor acetylhydrolase, isoform 1b, subunit 1) [NCBI Gene 18472] {aka LIS-1, Lis1, MMS10-U, Mdsh, Ms10u, Pafaha}, Krt5 (keratin 5) [NCBI Gene 369017] {aka CK-5, K5, Krt2-5, Sak57}, Ift81 (intraflagellar transport 81) [NCBI Gene 12589] {aka Cdv-1, Cdv-1r, Cdv1}, Foxc1 (forkhead box C1) [NCBI Gene 17300] {aka FREAC3, Fkh1, Mf1, Mf4, ch, fkh-1}, Mkks (McKusick-Kaufman syndrome) [NCBI Gene 59030] {aka 1300013E18Rik, Bbs6}, Afdn (afadin, adherens junction formation factor) [NCBI Gene 17356] {aka 5033403D15Rik, AF6, Af-6, Afadin, Gm314, I-afadin}, DPY19L2 (dpy-19 like 2) [NCBI Gene 283417] {aka SPATA34, SPGF9}, Ift27 (intraflagellar transport 27) [NCBI Gene 67042] {aka 2600013G09Rik, Rabl4}, Ccdc181 (coiled-coil domain containing 181) [NCBI Gene 74895] {aka 4930455F23Rik}, Tcp1 (t-complex protein 1) [NCBI Gene 21454] {aka CCT, Cct1, Ccta, TRic, Tcp-1, Tp63}, Lmna (lamin A) [NCBI Gene 16905] {aka Dhe}, H1f2 (H1.2 linker histone, cluster member) [NCBI Gene 50708] {aka 0610008C09Rik, H1-2, H1.2, H1c, H1var1, His1a}, Krt5 (keratin 5) [NCBI Gene 110308] {aka 3300001P10Rik, CK5, K5, Krt2-5, Tfip8}, CTTN (cortactin) [NCBI Gene 2017] {aka EMS1}, ZPBP (zona pellucida binding protein) [NCBI Gene 11055] {aka SPGF66, ZPBP1}, Cep131 (centrosomal protein 131) [NCBI Gene 12009] {aka AZ1, Azi, Azi1}, Spem1 (spermatid maturation 1) [NCBI Gene 74288] {aka 1700095G12Rik}, Ift88 (intraflagellar transport 88) [NCBI Gene 305918] {aka Ttc10}, Wasl (WASP like actin nucleation promoting factor) [NCBI Gene 73178] {aka 2900021I12Rik, 3110031I02Rik, N-WASP}, Mapre3 (microtubule-associated protein, RP/EB family, member 3) [NCBI Gene 100732] {aka EB2, EB3, EBF3, RP3}, Msn (moesin) [NCBI Gene 17698], Ubqln1 (ubiquilin 1) [NCBI Gene 56085] {aka 1110046H03Rik, 1810030E05Rik, D13Ertd372e, Da41, Dsk2, Plic-1}, Katnal2 (katanin p60 subunit A-like 2) [NCBI Gene 71206] {aka 3110023G01Rik, 4933439B08Rik}, Lmna (lamin A/C) [NCBI Gene 60374], PMFBP1 (polyamine modulated factor 1 binding protein 1) [NCBI Gene 83449] {aka SPGF31, STAP}, Cwc15 (CWC15 spliceosome-associated protein) [NCBI Gene 66070] {aka 0610040D20Rik, 2900052N06Rik, Ed1, c11orf5, mED1}, Nup62 (nucleoporin 62) [NCBI Gene 18226] {aka D7Ertd649e, Nupc1, p62}, Cntrob (centrobin, centrosomal BRCA2 interacting protein) [NCBI Gene 216846] {aka 9830165K03Rik, Lip8, Nip2}, Sun3 (Sad1 and UNC84 domain containing 3) [NCBI Gene 194974] {aka Sunc1}, Bbs5 (Bardet-Biedl syndrome 5) [NCBI Gene 72569] {aka 1700049I01Rik, 2700023J09Rik}, Spag4 (sperm associated antigen 4) [NCBI Gene 245865] {aka 1700041K21Rik, MNCb-0953, Sun4, mKIAA4118}, Ift172 (intraflagellar transport 172) [NCBI Gene 67661] {aka 4930553F24Rik, Slb, avc1, wim}, Rac1 (Rac family small GTPase 1) [NCBI Gene 19353] {aka D5Ertd559e} - **Diseases:** Didelphys sp (MESH:D000093642), MOLECULAR MECHANISMS (MESH:D041781), genetic defects (MESH:D030342), SPERMATOGENESIS (MESH:C536875), FORMATION (MESH:D058426), acephalic spermatozoa syndrome (OMIM:617187), head abnormalities (MESH:D006258), XLV-15 (MESH:D012559), male infertility (MESH:D007248), acrosomal defects (OMIM:102530), sperm abnormalities (MESH:C567467), MAMMALIAN (MESH:C000655084), MS (MESH:D009103), XL-28 (MESH:D000080345), infertility (MESH:D007246), immunodeficient (MESH:D007153), GRANTS (MESH:C537293), globozoospermia (MESH:D000072660) - **Species:** Cercopithecidae (monkey, family) [taxon 9527], Peromyscus (genus) [taxon 10040], Homo sapiens (human, species) [taxon 9606], Homo sp. (species) [taxon 2813599], Saccharomyces cerevisiae (baker's yeast, species) [taxon 4932], Selachii (sharks, infraclass) [taxon 119203], Testudines (anapsid reptiles, order) [taxon 8459], Vombatidae gen. sp. (wombat, species) [taxon 9339], Macaca (macaque, genus) [taxon 9539], Sus scrofa (pig, species) [taxon 9823], Cricetus cricetus (black-bellied hamster, species) [taxon 10034], Bufo bufo (common European toad, species) [taxon 8384], Pristis pristis (common sawfish, species) [taxon 1577861], Rajiformes (skates, order) [taxon 7858], Neoceratodus forsteri (Australian lungfish, species) [taxon 7892], Hyla arborea (common tree frog, species) [taxon 121869], Globicephala sp. (species) [taxon 77568], Rattus norvegicus (brown rat, species) [taxon 10116], Chiroptera (bats, order) [taxon 9397], Echidna (genus) [taxon 189497], Chimaera sp. (species) [taxon 30516], Sciurus vulgaris (Eurasian red squirrel, species) [taxon 55149], Macaca sylvanus (Barbary ape, species) [taxon 9546], Elasmobranchii (elasmobranchs, subclass) [taxon 7778], Testudo sp. (species) [taxon 2304327], Cricetulus griseus (Chinese hamster, species) [taxon 10029], Saimiri sciureus (common squirrel monkey, species) [taxon 9521], Gymnophiona (caecilians, order) [taxon 8445], Danio rerio (leopard danio, species) [taxon 7955], Chaetophractus villosus (large hairy armadillo, species) [taxon 29080], Gallus sp. (species) [taxon 9036], Equus caballus (domestic horse, species) [taxon 9796], Cavia sp. (species) [taxon 10143], Lepidosauria (lepidosaurs, class) [taxon 8504], Microtus arvalis (common vole, species) [taxon 47230], Dromiciops gliroides (monito del monte, species) [taxon 33562], Dipnomorpha (dipnoans, clade) [taxon 7878], Cricetinae (hamsters, subfamily) [taxon 10026], Passer domesticus (Haussperling, species) [taxon 48849], Didelphidae (American opossums, family) [taxon 9265], Cervus elaphus (red deer, species) [taxon 9860], Meles sp. (species) [taxon 30545], Sphenodon punctatus (hatteria, species) [taxon 8508], Salamandra salamandra (European fire salamander, species) [taxon 57571], Lemur sp. (species) [taxon 69491], Caudata (salamanders, order) [taxon 8293], Salmo sp. (species) [taxon 8031], Cetacea (cetaceans, infraorder) [taxon 9721], Cervidae (deer, family) [taxon 9850], Chaetognatha (arrow worms, phylum) [taxon 10229], Sphenodontia (beaked reptiles, order) [taxon 8505], Drosophila melanogaster (fruit fly, species) [taxon 7227], Phascolarctos cinereus (koala, species) [taxon 38626], Mus musculus (house mouse, species) [taxon 10090], Oryctolagus cuniculus (domestic rabbit, species) [taxon 9986], Anthozoa (anthozoans, class) [taxon 6101], Testudinidae (tortoises, family) [taxon 8487], Ovis aries (domestic sheep, species) [taxon 9940], Squamata (squamates, order) [taxon 8509], Canis lupus familiaris (dog, subspecies) [taxon 9615] ## Figures 11 figures with captions in the complete paper: https://tomesphere.com/paper/PMC8812575/full.md --- Source: https://tomesphere.com/paper/PMC8812575