Major histocompatibility complex similarity and sexual selection: different does not always mean attractive

Females that mate multiply have the possibility to exert postcopulatory choice and select more compatible sperm to fertilize eggs. Prior work suggests that dissimilarity in major histocompatibility complex (MHC) plays an important role in determining genetic compatibility between partners. Favouring a partner with dissimilar MHC alleles would result in offspring with high MHC diversity and therefore with enhanced survival thanks to increased resistance to pathogens and parasites. The high variability of MHC genes may further allow discrimination against the sperm from related males, reducing offspring homozygosity and inbreeding risk. Despite the large body of work conducted at precopulatory level, the role of MHC similarity between partners at postcopulatory level has been rarely investigated. We used an internal fertilizing fish with high level of multiple matings (Poecilia reticulata) to study whether MHC similarity plays a role in determining the outcome of fertilization when sperm from two males compete for the same set of eggs. We also controlled for genomewide similarity by determining similarity at 10 microsatellite loci. Contrary to prediction, we found that the more MHC‐similar male sired more offspring while similarity at the microsatellite loci did not predict the outcome of sperm competition. Our results suggest that MHC discrimination may be involved in avoidance of hybridization or outbreeding rather than inbreeding avoidance. This, coupled with similar findings in salmon, suggests that the preference for MHC‐dissimilar mates is far from being unanimous and that pre‐ and postcopulatory episodes of sexual selection can indeed act in opposite directions.     

Experimental evidence that high levels of inbreeding depress sperm competitiveness

The effects of inbreeding on sperm quantity and quality are among the most dramatic examples of inbreeding depression. The extent to which inbreeding depression results in decreased fertilization success of a male’s sperm, however, remains largely unknown. This task is made more difficult by the fact that other factors, such as cryptic female choice, male sperm allocation and mating order, can also drive patterns of paternity. Here, we use artificial insemination to eliminate these extraneous sources of variation and to measure the effects of inbreeding on the competitiveness of a male’s sperm. We simultaneously inseminated female guppies (Poecilia reticulata) with equal amounts of sperm from an outbred (f = 0) male and either a highly (f = 0.59) or a moderately inbred (f = 0.25) male. Highly inbred males sired significantly fewer offspring than outbred males, but share of paternity did not differ between moderately inbred and outbred males. These findings therefore confirm that severe inbreeding can impair the competitiveness of sperm, but suggest that in the focal population inbreeding at order of a brother–sister mating does not reduce a male’s sperm competitiveness.     

Female preference for male lateral projection area in <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

The lateral projection area (LPA) hypothesis was investigated by presenting female guppies, Poecilia reticulata with a series of paired males in three experiments. One trait was held constant in each experiment (dorsal fin size in expt I, body size in expt II, and total LPA in expt III) while body: dorsal fin ratio varied. Females spent more time near dummies of greater body and dorsal fin size in experiments I and II. The preference functions based on the first two sets of stimuli showed a similar pattern: the greater the LPA difference between paired dummies, the stronger the preference for the larger of the two. However, in the third experiment, neither fin size, body size, nor any particular dorsal fin + body size combination was preferred. These findings support the LPA hypothesis suggesting that increased LPA is more stimulating to sexually receptive females, and that females consequently prefer larger males. Preference for increased male dorsal fin size/LPA in both P. latipinna and P. mexicana as demonstrated in previous studies, and the more distantly related P. reticulata (shown here) is consistent with common ancestry of the sensory/neural systems in females of all three species and perhaps the entire Poeciliid family.     

Determinants of sperm precedence in a noctuid moth Heliothis virescens: a role for male age

1. Females of the noctuid moth Heliothis virescens F. mate more than once. Thus, sperm from two or more males normally compete for fertilisations within the female reproductive tract. The eggs are typically fertilised by sperm from only one male, either the female's last mate or an earlier mate. Twice‐mated females store only one ejaculate's worth of fertilising sperm (eupyrene) but nearly two ejaculates' worth of a nonfertilising sperm morph (apyrene), which is thought to play a role in sperm competition. 2. The mechanism of sperm use in H. virescens was investigated by examining factors that vary with paternity, which was assigned based on allozyme variation. The factors included male and female body masses and ages, male genital characters, the size of the sperm package, and the number of sperm stored by the female. 3. One male typically gained sperm precedence; this was nearly twice as likely to be the second male as it was to be the first. Two factors were found to vary significantly with paternity: female mass and male age. The second male to mate was more likely to gain sperm precedence if the female was larger and if the male was older than the female's first mate. 4. The significance of male age and female mass to several hypothetical models of the mechanism of sperm use is discussed.     

Determinants of male ejaculate investment in the cotton bollworm Helicoverpa armigera: mating history,female body size and male age

Many studies demonstrate that ejaculate size may be influenced by male condition, female quality and the risk or intensity of sperm competition. In the present study, the effect of male and female conditions, male mating history and female mating status on ejaculate sperm numbers in the polyandrous moth Helicoverpa armigera is examined. A large variation in ejaculate size is found and, although female body size and male age influence ejaculate size, female age and copula duration do not. Both male and female mating histories have significant effects on ejaculate sperm numbers. Males reduce ejaculate expenditure in successive matings but deliver significantly more apyrene and eupyrene sperm to nonvirgin than to virgin females.     

Factors Affecting Sperm Quality Before and After Mating of Calopterygid Damselflies

Damselflies (Odonata: Zygoptera) have a more complex sperm transfer system than other internally ejaculating insects. Males translocate sperm from the internal reproductive organs to the specific sperm vesicles, a small cavity on the body surface, and then transfer them into the female. To examine how the additional steps of sperm transfer contribute to decreases in sperm quality, we assessed sperm viability (the proportion of live sperm) at each stage of mating and after different storage times in male and female reproductive organs in two damselfly species, Mnais pruinosa and Calopteryx cornelia. Viability of stored sperm in females was lower than that of male stores even just after copulation. Male sperm vesicles were not equipped to maintain sperm quality for longer periods than the internal reproductive organs. However, the sperm vesicles were only used for short-term storage; therefore, this process appeared unlikely to reduce sperm viability when transferred to the female. Males remove rival sperm prior to transfer of their own ejaculate using a peculiar-shaped aedeagus, but sperm removal by males is not always complete. Thus, dilution occurs between newly received sperm and aged sperm already stored in the female, causing lower viability of sperm inside the female than that of sperm transferred by males. If females do not remate, sperm viability gradually decreases with the duration of storage. Frequent mating of females may therefore contribute to the maintenance of high sperm quality.     

Sperm displacement behavior of the cuttlefish Sepia esculenta (Cephalopoda: Sepiidae)

Sperm displacement behavior of cuttlefish (Sepia esculenta) was observed in a tank. Before ejaculation, male cuttlefish used their arms III to scrape out sperm masses attached to the buccal membranes of females. The removed sperm mass debris was directly visible and countable. Active sperm were present within the removed sperm debris, implying that the aim of this behavior is to remove competing male sperm. However, many sperm masses remained on the female buccal membrane even after the removal behavior, showing that sperm removal in S. esculenta is incomplete. The duration of sperm removal (an indicator of male investment in that process) was unaffected by the body sizes of mated pair, the duration of spermatangia placement at the current mating (for the hypothesis that the sperm removal serves to creat attachment space of spermatophores), or the estimated amount of sperm masses deposited from previous matings. Moreover, male S. esculenta performed sperm removal regardless of whether the last male to mate with the partner was himself, suggesting males remove not only the sperm of rivals but also their own. Although the number of removed sperm masses increased with the time spent on removal of sperm, male cuttlefish may shorten the duration of sperm removal to avoid the risk of mating interruption. We conclude that this time restriction would likely influence the degree of partial sperm removal in S. esculenta. A digital video image relating to the article is available at .This revised version was published online in April 2005 with corrections in the abstract.     

Enhanced activities of reproductive system in male rat treated with male silkworm pupae extract

A male silkworm pupae extract with herbal mixtures such as Rubus coreanus Miquel, Chinese matrimony vine Acanthopanax senticosus and tocopherol can be effectively used to recover or boost stamina. In this study, the effect of the male silkworm pupae extract on the reproductive system of Sprague–Dawley male rats was investigated. No clinical symptoms, and no dying or dead animals were found among experimental male silkworm larvae extract, male silkworm pupae extract and control rat groups during the study. No significant differences were found in body weights, feed or water consumption, or macroscopic examination among the three groups. No lesion was found upon necropsy. However, sperm in the group treated with male silkworm pupae extract were significantly more active than sperm in the control group. Sperm counts in both male groups treated with male silkworm larvae extract and male silkworm pupae extract were significantly higher than sperm counts in the control group. Analysis of phospholipid–hydroperoxide glutathione peroxidase (PHGPx) gene expression showed that the male silkworm pupae extract increased expression in testes by 26.7%. This study shows that the male silkworm pupae extract has potential to be used to enhance the function of the human reproductive system.     

Ultrastructure of spermatozoa of orsolobidae (Haplogynae,Araneae) with implications on the evolution of sperm transfer forms in Dysderoidea

Haplogynae are highly diverse with respect to the primary male genital system and sperm characteristics. Additionally, all sperm transfer forms (STF) known for spiders are present. Besides individually transferred sperm (cleistospermia), sperm are transferred as conjugates, both primary (synspermia) and secondary sperm conjugates (coenospermia, rouleaux) occur. Nevertheless, the ultrastructure of spermatozoa and STF are described for few Haplogynae and often only one representative species was studied, resulting in a superficial insight in the evolution of these traits. To elucidate the evolution of STF within Haplogynae we investigated representatives of four genera of the dysderoid family Orsolobidae. Our data show the presence of synspermia (Orsolobus, Osornolobus, Hickmanolobus, and Tasmanoonops) and also cleistospermia (Osornolobus). The occurrence of different STF within one family or even genus has not been described for any other spider taxon so far. Moreover, the synspermia of species of Tasmanoonops and Hickmanolobus were not covered by a secretion sheath suggesting a previously unknown strategy of transferring sperm that is possibly related to sperm residency time or female triggered processes after copulation. Based on serial ultrathin sectioning and subsequent 3D‐reconstruction, we obtained detailed measurements revealing remarkable size differences of STF. To evaluate the previously suggested correlation with the most distal region of the spermophor inside the embolus (intromittent part of the copulatory organ) we measured the diameter of the spermophor using micro‐computed X‐ray tomography data to obtain corresponding morphometric parameters. Based on these data only two species show similarity in STF and spermophor diameter. J. Morphol. 275:1238–1257, 2014. © 2014 Wiley Periodicals, Inc.     

Gamete number and size correlate with adult size in the egg parasitoid Trichogramma euproctidis

In parasitoids, the size of the adult is influenced by the size and quality of the host in which it develops. Body size is generally positively correlated with several adult fitness proxies (fecundity, longevity, and mating capacity). The initial resources available to an individual can influence gamete production (sperm and oocytes), and the number and quality of gametes produced directly influence the expected fitness of both males and females. Gamete production in relation to adult body size was quantified in Trichogramma euproctidis (Girault) (Hymenoptera: Trichogrammatidae), a short‐lived egg parasitoid of lepidopteran species. To avoid host quality variation, male and female parasitoids of different body sizes were produced using superparasitism by allowing mated and virgin female parasitoids to oviposit on Trichoplusia ni Hübner (Lepidoptera: Noctuidae) eggs. Seminal vesicles and ovaries of their offspring were dissected to count oocytes and to measure sperm length and oocytes volume. Tibia length was also measured to estimate body size. The number of oocytes, volume of oocytes, maternal investment index [= (number of oocytes × mean volume of oocytes)/10 000] and sperm length were all significantly positively correlated to body size. These results show that initial resources acquired during larval stage induce phenotypic plasticity in gamete production in both male and female T. euproctidis. Whereas number of sperm and oocytes can influence the fitness of males and females through increased mating capacity and fecundity, variation in gamete size (sperm length and oocyte volume) could also affect the fitness of an individual through sperm and larval competition.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

Directional postcopulatory sexual selection is associated with female sperm storage in Trinidadian guppies

Female sperm storage (FSS) is taxonomically widespread and often associated with intense sperm competition, yet its consequences on postcopulatory sexual selection (PCSS) are poorly known. Theory predicts that FSS will reduce the strength of PCSS, because sperm characteristics favored before and after FSS may be traded‐off, and opportunities for nondirectional PCSS should increase. We explored these questions in the guppy (Poecilia reticulata), by allowing females to mate multiply and by comparing the paternity pattern in two successive broods. Contrary to predictions, the variance in male fertilization success increased after FSS, driven by a change in male paternity share across broods. This change was positively associated with sperm velocity (measured before FSS) but not with the duration of FSS, indirectly suggesting that faster sperm were better in entering female storage organs, rather than in persisting within them. Other male traits, such as male size and orange color, heterozygosity, and relatedness to the female, did not influence paternity after FSS. These results indicate that processes associated with FSS tend to reinforce the strength of PCSS in guppies, rather than weaken it. Further work is necessary to test whether this pattern changes in case of more prolonged FSS.     

A new species of Ceriodaphnia from Uzbekistan and Kazakhstan

A Ceriodaphnia has been found inWaterbodies of the plains and foothills of Uzbekistan and southern Kazakhstan that strongly resembles C. reticulata (Jurine, 1820). The female of this new species, C. turkestanica, differs from C. reticulata because it lacks a fenestra (nucal organ) in the cervical notch of the adult, has a rostral pore encircled by a ridge of cuticle rather than opening on the flat surface of a polygon anterior to the antennules, and has fewer and finer spines punctuating the row of fine spinules along the inner margin of the posterior carapace. The locule of the ephippium lacks the regular, small, rounded protuberances seen in the locule of C. reticulata. The male of C. turkestanica also exhibits the lack of a cervical fenestra, an encircled rostral pore, and variable, fine spines along the posterior carapace inner margin. In addition, it has a much shorter terminal seta on the antennule, and increasing lengths of the spinose setae on endites 1–3 of trunk limb I rather than the second being the longest as in C. reticulata. Because of similarities between these species, we suggest that C. turkestanica either is derived from C. reticulata or that the two share a common ancestor.     

The effect of female presence on spermiation, and of male sexual activity on ‘ready’ sperm in the male guppy

This study examines the effect of female presence on the amount of ‘strippable’ sperm, and the relationship between male sexual activity and the amount of ‘ready’ sperm in the male guppy Poecilia reticulata. The amount of strippable sperm was greater in males paired for one week with females than in isolated males. We separated these ‘paired’ males from females by a clear partition and thus exposed males to visual and possibly olfactory stimuli. In the second experiment, control males spent more time attending females and performed more sexual acts during a 15-min test period than males previously stripped of spermatophores. These results indicate that females play an important role in stimulating sperm availability in males and that the amount of ready sperm influences male sexual behaviour. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Sperm morphology in the black coral Cirrhipathes sp. (Anthozoa, Antipatharia)

Abstract. Male polyps of the antipatharian Cirrhipathes sp., collected along the coral reef of Siladen Island (Sulawesi, Indonesia), were studied in order to gain an insight into the reproductive biology. Spermatocysts (maximum size 120 μm) are located within the primary gametogenic mesenteries and are separated by mesenteric cell cytoplasmic extensions. Sperm, maturing along radial rows, have a fairly round shape and contain a series of electron-dense vesicles in the apical nuclear region. A single mitochondrion flanks the nucleus. A peculiar cup-like electron-dense body, edged with regularly spaced electron-dense granules, is interposed between the nucleus and the tail, and delimits a central region that includes two centrioles. Cross-sections of the cup-like body reveal that the distal centriole has a pericentriolar system, consisting of nine arms arranged in a radial pattern. Each arm branches into three processes that are connected to the electron-dense granules. Indirect evidence of spawning is derived from the accumulation of sperm in the gastric cavity. This process takes place through the lysis of the cells bordering the mesenteries. Intact cells of this bordering layer appear to be involved in the phagocytosis of non-expelled gametes.     

Nereis succinea nuptial behavior: Does size matter?

Summary

Although Darwin noted that size may affect sexual selection, the effect of size on reproduction is controversial. Mature Nereis succinea, a polychaete that mates in pheromone-mediated twilight nuptial swarms, varies greatly in size, ranging more than 10-fold in body weight from 30 mg to >300 mg. Swim speed of swarming male worms increases with worm size, with the fastest males swimming more than twice the speed of the slowest. The female-produced spawning pheromone, nereithione, stimulates both swimming speed and spawning in males at concentrations of 10^?6 M and above, and lower concentrations cause significant activation of swimming. Individual worms can be stimulated to release sperm up to 40 or more times in a single experimental session. Larger worms release cumulatively more sperm than smaller ones, resulting in significant loss of body mass from repetitive spawning activated by nereithione. Size may enhance mating success of male N. succinea due to encountering more females as a result of faster swimming speed and due to the higher sperm density and number of spawning responses of large animals.     

Marked sperm dimorphism in the ground beetle Scarites terricola: a novel type of insect sperm polymorphism

Sperm polymorphism describes the phenomenon of male ejaculates containing two or more distinct types of sperm. In insects, four types of sperm polymorphism are recognized in species from the orders Diptera, Hemiptera, Hymenoptera, Lepidoptera and Coleoptera. The present study describes dimorphic sperm of the ground beetle Scarites terricola (Coleoptera: Carabidae) as a novel type of sperm polymorphism in insects. Sperm from the spermatophore and male seminal vesicles are examined at the light‐microscopic level, and both display marked dimorphism. One type has sperm formed into bundles, in which the head of numerous spermatozoa are ‘glued’ together, with tails free‐moving. The other type are free as single spermatozoa and have a disproportionately large‐sized head and an elongated tail. Both types are motile in Ringer's solution. The adaptive and phylogenetic importance of these findings is discussed.     

External structures used during attachment and sperm transfer in tubificids (Annelida,Oligochaeta)

The genital region of seven species of Tubificidae has been studied by SEM (Scanning Electron Microscopy). The form and the position of penial and spermathecal chaetae, male and spermathecal pores and other special structures have been examined. Peristodrilus montanus shows a special system to hold the partner: the penial chaetae anchor in an elaborated structure of the body wall formed between the spermathecal pores, the `anchorage bridge'. Protuberodrilus tourenqui has a long glandular porophore with the male pores at the tip, allowing contact with the spermathecal pores which are located in deep, close to the mid-ventral line of the body. The grooved and strongly curved penial chaetae of Rhyacodrilus falciformis seem to be used both for attachment and for sperm transfer, entering into the lateral spermathecal pores. The embrace of the partners, as suggested by observations on Psammoryctides barbatus, Potamothrix bavaricus, Potamothrix hammoniensis and Potamothrix heuscheri, seems to be another important mechanism to fix contact between male and spermathecal pores and allow sperm transfer. The spermathecal chaetae could be interpreted as piercing chaetae with a chemical or mechanical stimulating role. Sensitive cilia near the penial chaetae seem to be used by the three rhyacodrilines studied to find the correct anchorage place. There is a great variety of structures which appear to be used for attachment and sperm transfer in tubificids, and consequently their role in the evolution of the whole family may be profound.     

Fine structural observations of sperm resorption in the seminal vesicle of a marine snail,Littorina scutulata (Gould, 1849)

Summary Waste sperm and spermatozeugmata in the seminal vesicle of Littorina scutulata are phagocytised either by cell buds (large vesicles given off from the epithelial cells) or by the epithelial cells themselves. Cell buds containing sperm, are in turn engulfed by epithelial cells. In both cases, heterophagic vacuoles are formed inside the cell and subsequently the vacuoles are fused with primary lysosomes or lysosomal derivatives to become secondary lysosomes. Throughout this process the sperm are being digested. The second lysosome transforms further to telolysosome and finally to residual body when the sperm is completely digested.