MALE CONTEST COMPETITION AND THE COEVOLUTION OF WEAPONRY AND TESTES IN PINNIPEDS

Male reproductive success is influenced by competitive interactions during precopulatory and postcopulatory selective episodes. Consequently, males can gain reproductive advantages during precopulatory contest competition by investing in weaponry and during postcopulatory sperm competition by investing in ejaculates. However, recent theory predicts male expenditure on weaponry and ejaculates should be subject to a trade‐off, and should vary under increasing risk and intensity of sperm competition. Here, we provide the first comparative analysis of the prediction that expenditure on weaponry should be negatively associated with expenditure on testes mass. Specifically, we assess how sexual selection influences the evolution of primary and secondary sexual traits among pinnipeds (seals, sea lions, and walruses). Using recently developed comparative methods, we demonstrate that sexual selection promotes rapid divergence in body mass, sexual size dimorphism (SSD), and genital morphology. We then show that genital length appears to be positively associated with the strength of postcopulatory sexual selection. However, subsequent analyses reveal that both genital length and testes mass are negatively associated with investment in precopulatory weaponry. Thus, our results are congruent with recent theoretical predictions of contest‐based sperm competition models. We discuss the possible role of trade‐offs and allometry in influencing patterns of reproductive trait evolution in pinnipeds.     

THE ORIGIN OF THE SPINNING APPARATUS IN SPIDERS

• 1 Previous attempts to explain the evolution of spider silk have relied heavily on conjecture. The formulation of testable historical hypotheses to replace such speculation is discussed.
• 2 The importance of phylogenetic reconstructions and other historical hypotheses for use in generating and testing hypotheses concerning the evolution of specific adaptations is examined. Recent ideas on arachnid phylogeny are reviewed and their relevance to the problem of silk evolution in spiders is explored.
• 3 Evidence from the analysis of three historical problems (origin of spinnerets, origin of silk glands, original selective pressure favouring evolution of silk) is reviewed from three different frames of reference (in-group analysis, out-group analysis, convergence analysis). Several lines of evidence are found which suggest that silk use originated in spiders due to selective pressures associated with reproduction (specifically, the transfer of sperm or the protection of eggs).
• 4 The prevalence of segmental appendages retained for use in manipulating genital products in both arachnids and non-arachnid arthropods and the probable placement of spinnerets near the genital opening in ancestral spiders suggest that spinnerets represent modified gonopods.
• 5 The most primitive types of silk glands are retained in virtually all spiders, in part, for use in the construction of sperm webs and egg sacs. Similar silk glands are found near the genital opening in many male spiders and used in building a portion of the sperm web.
• 6 The silk of adult arthropods other than spiders is used largely in manipulating or protecting sex cells. If there are multiple functions, use in reproduction is typically one of them. Thus, there is evidence for strong selective pressure favouring the evolution of silk for use in reproduction.
• 7 Two hypotheses are proposed which are consistent with the conclusion that silk in spiders evolved for reproductive needs (the spermatophore-sperm web and egg sac hypotheses). Testable predictions of each hypothesis are proposed.

     

BOTH MALE AND FEMALE NOVEL TRAITS PROMOTE THE CORRELATED EVOLUTION OF GENITALIA BETWEEN THE SEXES IN AN ARTHROPOD

The correlated evolution of genitalia between sexes has been demonstrated in many taxa. However, it remains unclear whether female rather than male genitalia can play a key role in the correlated evolution of male and female genitalia. We conducted an extensive cross‐population analysis of the divergence patterns of genital structures, weights of whole genital organs, and the bodies of both sexes, and male genital length in a group of xystodesmid millipedes showing diverse genital morphologies. We demonstrate that the correlated evolution of male and female genitalia toward exaggerated states has occurred in the millipedes, which have evolved novel traits in both males (forceps‐like gonopods) and females (retractable bellows). Enlargement and elongation of forceps‐like gonopods may be advantageous in sperm competition, whereas enlargement and elongation of the bellows may facilitate acceptance/rejection of insemination for ensuring the female's fitness. These male and female genital parts have affected the correlated evolution in the opposite sex, resulting in diversification and exaggeration of genital morphology. Our study suggests that evolutionary novel traits in not only males but also in females could play an important role in the correlated evolution of genitalia between the sexes.     

One size fits all? Determinants of sperm transfer in a highly dimorphic orb‐web spider

The evolutionary significance of widespread hypo‐allometric scaling of genital traits in combination with rapid interspecific genital trait divergence has been of key interest to evolutionary biologists for many years and remains poorly understood. Here, we provide a detailed assessment of quantitative genital trait variation in males and females of the sexually highly dimorphic and cannibalistic orb‐weaving spider Argiope aurantia. We then test how this trait variation relates to sperm transfer success. In particular, we test specific predictions of the one‐size‐fits‐all and lock‐and‐key hypotheses for the evolution of genital characters. We use video‐taped staged matings in a controlled environment with subsequent morphological microdissections and sperm count analyses. We find little support for the prediction of the one‐size‐fits‐all hypothesis for the evolution of hypo‐allometric scaling of genital traits, namely that intermediate trait dimensions confer highest sperm transfer success. Likewise, our findings do not support the prediction of the lock‐and‐key hypothesis that a tight fit of male and female genital traits mediates highest sperm transfer success. We do, however, detect directional effects of a number of male and female genital characters on sperm transfer, suggesting that genital trait dimensions are commonly under selection in nature. Importantly, even though females are much larger than males, spermatheca size limits the number of sperm transferred, contradicting a previous hypothesis about the evolutionary consequences of genital size dimorphism in extremely size‐dimorphic taxa. We also find strong positive effects of male body size and copulation duration on the probability of sperm transfer and the number of sperm transferred, with implications for the evolution of extreme sexual size dimorphism and sexual cannibalism in orb weavers.     

THE CENTRAL ROLE OF THE GENITAL DUCT IN THE DEVELOPMENT AND REGENERATION OF THE GENITAL ORGANS IN THE SEA URCHIN

Formation and regeneration of the genital papilla in the sea urchin, Hemicentrotus pulcherrimus were studied to find out what factor(s) control expression of the secondary sexual characters in the papilla. Surgical removal of various regions of the genital organ and the blood vessels was performed separately or in various combinations. The proximal part of the genital duct or its rudiment (D2 region) plays an essential role in the formation or regeneration of the papilla, and in the establishment of its sexual characters as well, i.e., if the D2 region is removed, papilla formation or regeneration is completely suppressed; when the D2 region is left uninjured, the papilla develops and displays the sexual characters even in the absence of the gonad, the distal part of the duct, a blood supply to the D2 region or all three of them combined. It can be concluded that the proximal part of the genital duct carries the factor(s) for determination and expression of the secondary sexual characters in the genital papilla. Discussion has been made about what factor(s) can be considered to reside within the cells of the D2 region of the duct.     

Seasonal variation in genital and body size, sperm displacement ability, female mating rate, and male harassment in two calopterygid damselflies (Odonata: Calopterygidae)

Sperm competition is a pervasive force. One adaptation is the male ability to displace the rivals' sperm that females have stored from previous copulations. In the damselfly, Calopteryx haemorrhoidalis asturica , males with wider aedeagi displace more spermathecal sperm. The present study documents that the same mechanism operates in another damselfly, Hetaerina americana . However, this genital width in both species decreases along the season, but late-emerging females have more sperm displaced than early-emerging females. Because territorial males mated more and were larger in body and genital size than nonterritorial males, late-season females mated with considerably larger males with respect to female size and this produced higher sperm displacement. Assuming female benefits from storing sperm but that such benefit does not prevail if males displace sperm, it is predicted that, along the season, females will mate less and male harassment (in terms of male mating attempts and oviposition duration) will increase. These predictions were corroborated. In H. americana , it was also tested whether spermathecal sperm became less viable along the season. The results obtained did not corroborate this. This is the first evidence indicating that season affects sperm displacement ability and female mating frequency due to changes in male body and genital size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 815–829.     

Complex copulatory behavior and the proximate effect of genital and body size differences on mechanical reproductive isolation in the millipede genus Parafontaria

The role of species-specific genitalia in reproductive isolation is unclear. Males of the millipede genus Parafontaria use gonopods (modified eighth legs) charged with sperm from the genital openings of the second legs as intromittent organs. Males perform both preliminary and true intromission during mating. During preliminary intromission, a male attempts to insert his gonopods into the female genitalia before charging the gonopods with sperm. If this intromission is completed, it is followed by the ejaculation of sperm to the gonopods and true intromission for insemination. In two sympatric species of Parafontaria that lack effective precopulatory isolation, copulation was terminated without insemination because of preliminary intromission failure caused by mismatched genital and body sizes. Thus, mechanical isolation between these sympatric species resulted from morphological differentiation mediated by the obligatory preliminary intromission. These findings demonstrate the proximate importance of genital and body size differences for reproductive isolation within this genus of millipede.     

Expression of scatter factor/hepatocyte growth factor is regionally correlated with the initiation of sperm motility in murine male genital tract: Is scatter factor/hepatocyte growth factor involved in initiation of sperm motility?

Based upon findings that the scatter factor/hepatocyte growth factor (SF/HGF) has strong mitogenic and motogenic properties, and that the sperm cell acquires its fertilizing capacity and motility in the distal parts of mammalian epididymis, the present study was conducted to investigate the role of SF/HGF in initiation of sperm cell motility. This was investigated by determining the expression of SF/HGF in various regions of the murine male genital tract by scatter and cell tracking assays using MDCK epithelial cells, Western blot procedure, and the immunohistochemical procedure using paraffin sections of various regions of the male genital tract. The findings from all these assays indicate that SF/HGF is differentially expressed in various parts of the male genital tract with slight or no expression in the testes, caput epididymis, and vas deferens, and with the highest expression in cauda and corpus (distal) epididymis followed by expression in the corpus (proximal) epididymis. This region-specific SF/HGF expression pattern coincides with the pattern of acquiring the fertilizing capacity and motility by the sperm cell during its transit through the male genital tract. However, wherever SF/HGF was expressed in the male genital tract, its molecular weight was slightly higher (Mr, 82 kD), compared to the SF/HGF expressed in various other somatic tissues (Mr, 78 kD), indicating that the genital tract SF/HGF may be a different molecular species that shares some immunoreactive epitopes with the somatic cell SF/HGF. Incubation of immotile sperm from caput epididymis with the purified human placental SF/HGF of 78 kD initiated motility in 5–15% of sperm population. These results strongly suggest that the SF/HGF-like activity is expressed in the male genital tract in a region-specific manner, and this activity may have a role in initiation of sperm motility acquired during its transit through the epididymis in mammals. © 1994 Wiley-Liss, Inc.     

Morphology and histology of male and female reproductive systems in the inseminating species Scoloplax distolothrix (Ostariophysi: Siluriformes: Scoloplacidae)

The morphology and histology of male and female reproductive systems were examined in Scoloplax distolothrix. Internal insemination was documented in this species by the presence of sperm within the ovaries. Mature males and females have elongated genital papillae, exhibiting a tubular shape in males and a plain heart‐shape with two median protuberances in females. The testes are two elongated structures that converge ventrally, under the intestine, towards the genital papilla. They are joined at the caudal end, forming an ovoid single chamber for sperm storage. Secretory regions were not observed. In the lumen of the testicular tubules, spermatozoa can be tightly packed along their lengths, but do not constitute a spermatozeugmata. The lumen of the sperm storage chamber and spermatic duct are filled with free spermatozoa without the accompanying secretions. The ovaries are bird‐wing shaped, saccular structures that converge ventrally under the intestine, towards the genital papilla. They are joined at the caudal end, forming a tubular chamber possibly destined for oocyte storage. An oviduct with an irregular outline connects the chamber to the tubular region of the genital papilla. No distinct sperm storage structure was found in the ovaries. The unique male and female genital papillae suggest that these structures are associated with the reproductive mode in scoloplacids, representing evidence for insemination. The occurrence of free spermatozoa, without the accompanying secretions and not arranged in a spermatozeugmata can be associated with the presence of a tubular male genital papilla for sperm transfer to the female genital tract. This reinforces the idea that sperm packets are not necessary for all inseminating species. The male reproductive system in scoloplacids is very different from that in auchenipterids, a second catfish family with insemination, which indicates that the occurrence of insemination is not connected to the internal morphology of reproductive organs. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

THE POLYMERIZATION OF ACTIN: ITS ROLE IN THE GENERATION OF THE ACROSOMAL PROCESS OF CERTAIN ECHINODERM SPERM

When Asterias or Thyone sperm come in contact with egg jelly, a long process which in Thyone measures up to 90 µm in length is formed from the acrosomal region. This process can be generated in less than 30 s. Within this process is a bundle of microfilaments. Water extracts prepared from acetone powders of Asterias sperm contain a protein which binds rabbit skeletal muscle myosin forming a complex whose viscosity is reduced by ATP. Within this extract is a protein with the same molecular weight as muscle actin. It can be purified either by collecting the pellet produced after the addition of Mg⁺⁺ or by reextracting an acetone powder of actomyosin prepared by the addition of highly purified muscle myosin to the extract. The sperm actin can be polymerized and by electron microscopy the polymer is indistinguishable from muscle F-actin. The sperm actin was shown to be localized in the microfilaments in the acrosomal processes by: (a) heavy meromyosin binding in situ, (b) sodium dodecyl sulfate (SDS) gel electrophoresis of the isolated acrosomal processes and a comparison to gels of flagella which contain no band corresponding to the molecular weight of actin, and (c) SDS gel electrophoresis of the extract from isolated acrosomal caps. Since the precursor for the microfilaments in the unreacted sperm appears amorphous, we suspected that the force for the generation of the acrosomal process is brought about by the polymerization of the sperm actin. This supposition was confirmed, for when unreacted sperm were lysed with the detergent Triton X-100 and the state of the actin in the sperm extract was analyzed by centrifugation, we determined that at least 80% of the actin in the unreacted sperm was in the monomeric state.     

Spermatophore formation in the pseudoscorpion Chthonius ischnocheles (Chthoniidae)

The reproduction of pseudoscorpions involves indirect sperm transfer by means of spermatophores. The spermatophores are the product of the male genital atrium. A functional interpretation of spermatophore formation in Chthonius ischnocheles is based on evidence from (a) a morphological study of the genital atrium, the associated accessory glands and the musculature (b) males sectioned during spermatophore production (c) histochemical tests on the glands and their secretions (d) biochemical analyses of one gland (posterior dorsal) and its secretion (e) the behaviour of males during this process and (f) the structure of the spermatophore.
The anterior region of the genital atrium is concerned with the production of the sperm packet. The encysted sperm and the seminal fluid from the prostatic reservoir are encapsulated in a sperm packet by a secretion from two pairs of anterior glands. The posterior region of the genital atrium is responsible for the formation of the spermatophore stalk and its distal elaboration, the two lateral collars. These parts of the spermatophore arise from the fibroin secretion of the posterior dorsal gland; the shape of the spermatophore collars is correlated with their mould, the medial diverticula. In addition the lateral glands secrete a light oil which accumulates on a thickening of the spermatophore stalk proximal to the collars. This suggested that this secretion acts as a pheromone to attract females to the spermatophore since in this species males produce their spermatophores in the absence of females.     

THE REPRODUCTIVE ANATOMY OF PAROPEAS ACHATINACEUM AND A NEW CONCEPT OF PAROPEAS (PULMONATA: ACHATINOIDEA: SUBULINIDAE)

The majority of subulinid land snails are known only as shellsand generic groupings need to be revised on the basis of theirinternal anatomy. Paropeas achatinaceum (new combination) isa synanthropic subulinid that is widely distributed around theIndo-Pacific region. Features of the genital anatomy and radulademonstrate that Paropeas should be rejected as a subgenus ofProsopeas and be recognised as a distinct genus. The radulaof Paropeas is more typical of a subulinid than is that of Prosopeaswhich proves to be highly modified for a carnivorous diet. Thespermatophore of Paropeas is inserted directly into the oviductwhere it releases sperm through its porous tail before beingdigested in the bursa copulatrix. (Received 23 September 1992; accepted 11 November 1993)     

Male genital morphology and its influence on female mating preferences and paternity success in guppies

In internally fertilizing species male genitalia often show a higher degree of elaboration than required for simply transferring sperm to females. Among the hypotheses proposed to explain such diversity, sexual selection has received the most empirical support, with studies revealing that genital morphology can be targeted by both pre-and postcopulatory sexual selection. Until now, most studies have focused on these two episodes of selection independently. Here, we take an alternative approach by considering both components simultaneously in the livebearing fish, Poecilia reticulata. We allowed females to mate successively (and cooperatively) with two males and determined whether male genital length influenced the female's propensity to mate with a male (precopulatory selection, via female choice) and whether male genital size and shape predicted the relative paternity share of subsequent broods (postcopulatory selection, via sperm competition/cryptic female choice). We found no evidence that either episode of sexual selection targets male genital size or shape. These findings, in conjunction with our recent work exposing a role of genital morphology in mediating unsolicited (forced) matings in guppies, further supports our prior speculation that sexual conflict may be an important broker of genital evolution in this species.     

PREMATURE CHROMOSOME CONDENSATION OF THE KARYOGAMY-BLOCKED SPERM PRONUCLEUS IN THE FERTILIZATION OF FUCUS DISTICHUS (FUCALES,PHAEOPHYCEAE)1

Mitosis of egg and sperm pronuclei of Fucus distichus subsp. evanescens (C. Agardh)Powell was examined by fluorescence and electron microscopy when migration of the sperm pronucleus and, as a result, karyogamy were blocked by colchicine treatment after plasmogamy. Chromosome condensation was obsewed in both pronuclei Microspectrophotometric studies after staining the nuclei with mithramycin A clearly showed that DNA synthesis ocurred in the egg pronucleus but not in the sperm pronucleus. This means that chromosomes condensed prematurely in the sperm pronucleus (premature chromosome condensation). In some cases, the egg chromosomes became arranged on a metaphase plate, whereas the sperm chromosomes lay scattered near the egg pronucleus. Immuno fluorescence microscopy using anti-β-tubulin antibody confirmed that a normal spindle was formed at the egg pronucleus. A pair of centrioles existed at the two poles of this spindle. The sperm nuclear membrane disappeared, and microtubules radiated to the sperm chromosomes from one pole of the egg spindle.     

Sperm storage secretion of female cellar spiders (Pholcus phalangioides; Araneae): a gel-electrophoretic analysis

Glandular secretion in the genital cavity of female Pholcus phalangioides was removed and investigated gel-electrophoretically. Different staining techniques indicate that the secretion contains proteinaceous substances. Some protein fragments show a clear reaction to glyco- and lipoprotein staining procedures.
After copulation, female P. phalangioides keep the spermatozoa embedded in the secretion in the genital cavity. Since the spermatozoa are immobile, i.e. coiled and encapsulated, until shortly before oviposition, a nutritive function of the secretion during sperm storage is unlikely. The viscous quality of the material probably serves to retain the sperm mass in the sperm storage site. Lipoproteinaceous components are assumed to prevent desiccation of the sperm mass. Retaining the spermatozoa and maintaining them in a favourable environment are the most plausible functions of the secretion of female P. phalangioides. In spider species which possess receptacula seminis other functions can be attributed to sperm storage secretions.     

The evolution of asymmetric genitalia in spiders and insects

Asymmetries are a pervading phenomenon in otherwise bilaterally symmetric organisms and recent studies have highlighted their potential impact on our understanding of fundamental evolutionary processes like the evolution of development and the selection for morphological novelties caused by behavioural changes. One character system that is particularly promising in this respect is animal genitalia because (1) asymmetries in genitalia have evolved many times convergently, and (2) the taxonomic literature provides a tremendous amount of comparative data on these organs. This review is an attempt to focus attention on this promising but neglected topic by summarizing what we know about insect genital asymmetries, and by contrasting this with the situation in spiders, a group in which genital asymmetries are rare. In spiders, only four independent origins of genital asymmetry are known, two in Theridiidae (Tidarren/Echinotheridion, Asygyna) and two in Pholcidae (Metagonia, Kaliana). In insects, on the other hand, genital asymmetry is a widespread and common phenomenon. In some insect orders or superorders, genital asymmetry is in the groundplan (e.g. Dictyoptera, Embiidina, Phasmatodea), in others it has evolved multiple times convergently (e.g. Coleoptera, Diptera, Heteroptera, Lepidoptera). Surprisingly, the huge but widely scattered information has not been reviewed for over 70 years. We combine data from studies on taxonomy, mating behaviour, genital mechanics, and phylogeny, to explain why genital asymmetry is so common in insects but so rare in spiders. We identify further fundamental differences between spider and insect genital asymmetries: (1) in most spiders, the direction of asymmetry is random, in most insects it is fixed; (2) in most spiders, asymmetry evolved first (or only) in the female while in insects genital asymmetry is overwhelmingly limited to the male. We thus propose that sexual selection has played a crucial role in the evolution of insect genital asymmetry, via a route that is accessible to insects but not to spiders. The centerpiece in this insect route to asymmetry is changes in mating position. Available evidence strongly suggests that the plesiomorphic neopteran mating position is a female-above position. Changes to male-dominated positions have occurred frequently, and some of the resulting positions require abdominal twisting, flexing, and asymmetric contact between male and female genitalia. Insects with their median unpaired sperm transfer organ may adopt a one-sided asymmetric position and still transfer the whole amount of sperm. Spiders with their paired sperm transfer organs can only mate in symmetrical or alternating two-sided positions without foregoing transfer of half of their sperm. We propose several hypotheses regarding the evolution of genital asymmetry. One explains morphological asymmetry as a mechanical compensation for evolutionary and behavioural changes of mating position. The morphological asymmetry per se is not advantageous, but rather the newly adopted mating position is. The second hypothesis predicts a split of functions between right and left sides. In contrast to the previous hypothesis, morphological asymmetry per se is advantageous. A third hypothesis evokes internal space constraints that favour asymmetric placement and morphology of internal organs and may secondarily affect the genitalia. Further hypotheses appear supported by a few exceptional cases only.     

SPERM COMPETITION AND THE EVOLUTION OF NONFERTILIZING SPERM IN MAMMALS

Nonfertilizing sperm with special morphologies have long been known to exist in invertebrates. Until recently, abnormal sperm in mammals were considered errors in production. Now, however, Baker and Bellis (1988, 1989) have proposed that mammalian sperm, like some invertebrate sperm, are polymorphic and adapted to a variety of nonfertilizing roles in sperm competition, including prevention of passage of sperm inseminated by another male. More specifically, their “kamikaze” sperm hypothesis proposes that deformed mammalian sperm are adapted to facilitate the formation and functioning of copulatory plugs (Baker and Bellis, 1988). Here I argue that most, maybe all, mammals are unlikely to produce nonfertilizing sperm. First, mammals might not be able to afford to evolve nonfertilizing sperm, given that a) fertilization is often unlikely despite the huge numbers of sperm produced; b) production of larger numbers of sperm is constrained, presumably because of metabolic costs, evidence for which includes the fact that in species in which sperm morphology and anatomy of the female reproductive tract increase the probability of fertilization, the numbers of sperm produced is lower than in others; and c) selection appears to act against the production of deformed sperm. Second, some of the evidence advanced for the existence of nonfertilizing sperm does not in fact support the idea. Third, accessory gland secretions are sufficient on their own to coagulate semen and produce fully functioning plugs; thus the male that used accessory gland secretions would be at a clear advantage over the male that diluted his fertilizing sperm with “kamikaze” sperm; and indeed, current evidence indicates selection on accessory glands, not sperm morphology, to enhance coagulation of semen. Fourth, predictions made on the basis of the “kamikaze” sperm hypothesis are not supported by quantitative comparisons of data from polyandrous and monandrous primates (i.e., those in which several males mate with a fertile female, and therefore in which sperm competition should be operating, and those in which only one male mates). Although sperm competition is almost certainly more intense in polyandrous genera than in monandrous genera (as indicated by, e.g., more frequent copulations and the production of more sperm per ejaculate from larger spermatogenic organs), polyandrous genera do not produce a greater proportion of deformed (i.e., nonfertilizing) sperm than do monandrous genera, or even necessarily a greater number of deformed sperm; nor a greater variety of sperm sizes—indeed they might produce fewer; nor fewer motile sperm (as might be expected if sperm are selected to stay behind and compete with sperm from subsequent males); and nor larger sperm (as might be expected if sperm are produced for functions other than to reach the egg). In sum, currently available evidence suggests that the function of all mammalian sperm is to fertilize, and that sperm competition in mammals occurs through scramble competition, not contest competition.     

THE EVOLUTION OF SPERM SIZE IN BIRDS

Sperm size varies enormously among species, but the reasons for this variation remain obscure. Since it has been suggested that swimming velocity increases with sperm length, earlier studies proposed longer (and therefore faster) sperm are advantageous under conditions of intense sperm competition. Nonetheless, previous work has been equivocal, perhaps because the intensity of sperm competition was measured indirectly. DNA profiling now provides a more direct measure of the number of offspring sired by extrapair males, and thus a more direct method of assessing the potential for sperm competition. Using a sample of 21 species of passerine birds for which DNA profiling data were available, we found a positive relation between sperm length and the degree of extrapair paternity. A path analysis, however, revealed that this relationship arises only indirectly through the positive relationship between the rate of extrapair paternity and length of sperm storage tubules (SSTs) in the female. As sperm length is correlated positively with SST length, an increase in the intensity of sperm competition leads to an increase in sperm length only through its effect on SST length. Why females vary SST length with the intensity of sperm competition is not clear, but one possibility is that it increases female control over how sperm are used in fertilization. Males, in turn, may respond on an evolutionary time scale to changes in SST size by increasing sperm length to prevent displacement from rival sperm. Previous theoretical analyses predicting that sperm size should decrease as sperm competition becomes more intense were not supported by our findings. We suggest that future models of sperm-size evolution consider not only the role of sperm competition, but also how female control and manipulation of ejaculates after insemination selects for different sperm morphologies.     

THE FINE STRUCTURE OF NUCLEI DURING SPERM MATURATION IN THE LOCUST

1. In the heads of maturing sperm of a locust the chromatin becomes arranged in a highly regular manner so as to produce many parallel lines about 60 A thick in longitudinal section and contiguous polygons in transverse section. 2. This configuration appears after fixation in osmium tetroxide, formaldehyde, or acetic acid; intermediate stages in its development are illustrated. 3. These electron micrographs are interpreted to mean that, during sperm maturation, the chromatin becomes formed into sheets and then into tubes running parallel with the long axis. 4. In the mature sperm head we have been unable so far to detect this structure. This may be because the chromatin becomes so compact during the shrinkage of the nucleus which occurs during formation of the mature sperm.     

Modification of genital kidney nephrons for sperm transport in a plethodontid salamander,Eurycea longicauda

Salamanders possess kidneys with two distinct regions: a caudal pelvic portion and cranial genital portion. Nephrons of the pelvic region are responsible for urine formation and transport. Nephrons of the genital region transport sperm from testes to Wolffian ducts; however, nephrons of the genital region possess all the same functional regions found in pelvic kidney nephrons that are involved with urine formation and transport (renal corpuscles, proximal tubules, distal tubules, and collecting ducts). Morphological similarities between pelvic and genital regions stimulated past researchers to hypothesize that nephrons of genital kidneys possess dual function; that is, sperm transport and urine formation/transport. Considering size of glomeruli is directly related to the total amount of blood plasma filtered into the Bowman's space, we tested the hypothesis that nephrons of genital kidneys have reduced urine formation function by comparing glomerular size between nephrons of pelvic and genital kidney regions in Eurycea longicauda with general histological techniques. Light microscopy analysis revealed that glomeruli of pelvic kidneys were significantly larger than those measured from genital kidneys. Transmission electron microscopy analysis also revealed modifications in genital kidney nephrons when compared to pelvic kidney nephrons that suggested a decrease in urine formation function in genital kidneys. Such modifications included a decrease in basal and lateral plasma membrane folding in genital kidney proximal and distal tubules compared to that of pelvic kidney proximal and distal tubules. Genital kidney proximal tubules were also ciliated, which was not observed in pelvic kidney proximal tubules. In conclusion, although structurally similar at the histological level, it appears that nephrons of genital kidneys have decreased urine formation function based on glomerular size comparison and nephron ultrastructure.