So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

In the Hamadryas baboon, males are substantially larger than females. A troop of baboons is subdivided into a number of ‘one-male groups’, consisting of one adult male and one or more females with their young. The male prevents any of ‘his’ females from moving too far from him. Kummer (1971) performed the following experiment. Two males, A and B, previously unknown to each other, were placed in a large enclosure. Male A was free to move about the enclosure, but male B was shut in a small cage, from which he could observe A but not interfere. A female, unknown to both males, was then placed in the enclosure. Within 20 minutes male A had persuaded the female to accept his ownership. Male B was then released into the open enclosure. Instead of challenging male A , B avoided any contact, accepting A’s ownership.

Evolution and the Theory of Games

General Themes: G.A. Parker, Sperm Competition and the Evolution of Ejaculates: Towards a Theory Base. A.P. Moller, Sperm Competition and Sexual Selection. W.G. Eberhard, Female Roles in Sperm Competition. J. Wright, Paternity and Paternal Care. Taxonomic Treatments: L.F. Delph and K. Havens, Pollen Competition in Flowering Plants. D.R. Levitan, Sperm Limitation, Gamete Competition and Sexual Selection in External Fertilizers. N.K. Michiels, Mating Conflicts and Sperm Competition in Simultaneous Hermaphrodites. B. Baur, Sperm Competition in Molluscs. M.A. Elgar, Sperm Competition and Sexual Selection in Spiders and Other Arachnids. L.W. Simmons and M.T. Siva-Jothy, Sperm Competition in Insects: Mechanisms and the Potential for Selection. C.W. Petersen and R.R. Warner, Sperm Competition in Fishes. T.R. Halliday, Sperm Competition in Amphibians. M. Olsson and T. Madsen, Sexual Selection and Sperm Competition in Reptiles. T.R. Birkhead, Sperm Competition in Birds: Mechanisms and Function. D.A. Taggart, W.G. Breed, P.D. Temple-Smith, A. Purvis, and G. Shimmin, Reproduction, Mating Strategies and Sperm Competition in Marsupials and Monotremes. M. Gomendio, A.H. Harcourt, and E.R.S. Roldan, Sperm Competition in Mammals. T.R. Birkhead and A.P. Moller, Sperm Competition, Sexual Selection and Different Routes to Fitness. Index.

Sperm competition and sexual selection

ION, GENERAL CONCEPTIONS, SELF-CONSCIOUSNESS, MENTAL INDIVIDUALITY. It would be very difficult for any one with even much more knowledge than I possess, to determine how far animals exhibit any traces of these high mental powers. This difficulty arises from the impossibility of judging what passes through the mind of an animal; and again, the fact that writers differ to a great extent in the meaning which they attribute to the above terms, causes a further difficulty. If one may judge from various articles which have been published lately, the greatest stress seems to be laid on the supposed entire absence in animals of the power of abstraction, or of forming general concepts. But when a dog sees another dog at a distance, it is often clear that he perceives that it is a dog in the abstract; for when he gets nearer his whole manner suddenly changes, if the other dog be a friend. A recent writer remarks, that in all such cases it is a pure assumption to assert that the mental act is not essentially of the same nature in the animal as in man. If either refers what he perceives with his senses to a mental concept, then so do both. (44. Mr. Hookham, in a letter to Prof. Max Muller, in the 'Birmingham News,' May, 1873.) When I say to my terrier, in an eager voice (and I have made the trial many times), "Hi, hi, where is it?" she at once takes it as a sign that something is to be hunted, and generally first looks quickly all around, and then rushes into the nearest thicket, to scent for any game, but finding nothing, she looks up into any neighbouring tree for a squirrel. Now do not these actions clearly shew that she had in her mind a general idea or concept that some animal is to be discovered and hunted? It may be freely admitted that no animal is self-conscious, if by this term it is implied, that he reflects on such points, as whence he comes or whither he will go, or what is life and death, and so forth. But how can we feel sure that an old dog with an excellent memory and some power of imagination, as shewn by his dreams, never reflects on his past pleasures or pains in the chase? And this would be a form of self-consciousness. On the other hand, as Buchner (45. 'Conferences sur la Theorie Darwinienne,' French translat. 1869, p. 132.) has remarked, how little can the hardworked wife of a degraded Australian savage, who uses very few abstract words, and cannot count above four, exert her self-consciousness, or reflect on the nature of her own existence. It is generally admitted, that the higher animals possess memory, attention, association, and even some imagination and reason. If these powers, which differ much in different animals, are capable of improvement, there seems no great improbability in more complex faculties, such as the higher forms of abstraction, and selfconsciousness, etc., having been evolved through the development and combination of the simpler ones. It has been urged against the views here maintained that it is impossible to say at what point in the ascending scale animals become capable of abstraction, etc.; but who can say at what age this occurs in our young children? We see at least that such powers

https://www.jstage.jst.go.jp/article/ase1887/22/257/22_257_458/_pdf

The descent of man and selection in relation to sex

The evolution of polyandry : multiple mating and female fitness in insects

Sperm are produced in astronomical numbers compared with eggs, and there is good evidence that sperm competition is the force behind the evolution of many tiny sperm. However, sperm production inevitably has costs. Recent research shows that male ejaculate expenditure is dynamic in both time and space, and that males are sensitive to risks of sperm competition and can vary ejaculate size accordingly. We focus on studies showing that males assess mating status and relative fecundity of females, and reveal that modulation of ejaculate investment by males can sometimes result in sperm limitation for females.

Sperm competition, male prudence and sperm-limited females

Fishes show one of the widest ranges of sperm competition intensity of any animal group. Here we present a comparative study whose aim is to investigate the effect of relative intensity of sperm competition on investment in spermatogenesis and the number and size of sperm produced. We find that both the gonadosomatic index (GSI5 (gonad weight/body weight) 3 100) and sperm numbers increase with intensity of sperm competition across species but that sperm length decreases. These new findings are consistent with a raffle-based mode of sperm competition in fishes. Most of these results (positive correlation of the GSI and sperm number with sperm competition intensity) concur with the predictions of current sperm competition the- ory. However, we also find that sperm longevity decreases with sperm length across species. Cur- rent models for continuous fertilization suggest that if length increases a sperm's speed but de- creases its longevity, sperm length should increase with sperm competition intensity, whereas models for instant fertilization suggest that sperm length should remain constant. The negative relationship found between sperm competition and sperm length therefore does not fit predictions of either model.

https://www.jstor.org/stable/10.1086/286031

Sperm competition in fishes: the evolution of testis size and ejaculate characteristics.

https://www.cell.com/current-biology/pdf/S0960-9822(03)00939-4.pdf

Spermatozoal traits and sperm competition in Atlantic salmon: relative sperm velocity is the primary determinant of fertilization success.

This paper investigates mechanisms of sperm competition by comparing reproductive characteristics across 74 butterfly species. Testis size scales with body size and, after controlling for this allometry, relative testis size increases with risk of sperm competition, as defined by female mating frequency. Both eupyrene (fertilizing) and apyrene (non-fertile) sperm lengths correlate positively with body size. After controlling for body size, relative eupyrene sperm lengths are greater in species where males experience higher risks of sperm competition. These results suggest that sperm competition in butterflies selects for increased investment in spermatogenesis, and specifically longer fertilizing sperm. Because longer sperm may be faster and more powerful, eupyrene sperm may therefore compete energetically, and are not selected to be minimally sized to maximize numbers for a purely raffle-based sperm competition mode. Apyrene sperm lengths are not affected directly by risk of encountering rival sperm. Instead, apyrene sperm show closer associations with body size which, if female tract morphometry correlates with body size, is consistent with the hypothesis that apyrene sperm retard female sexual receptivity by moving while in storage.

Associations between Body Size, Mating Pattern, Testis Size and Sperm Lengths across Butterflies

We develop the logic of assessment of sperm competition risk by individual males where the mechanism of sperm competition follows a 'loaded raffle' (first and second inseminates of a female have unequal prospects). Male roles (first or second to mate) are determined randomly. In model 1, males have no information about the risk associated with individual females and ejaculation strategy depends only on the probability, q, that females mate twice. Evolutionarily stable strategy (ESS) ejaculate expenditure increases linearly from zero with q, and reduces with increasing inequality between ejaculates, though the direction of the loading (which role is favoured) is unimportant. In model 2, males have perfect information and can identify each of three risk states: females that will (1) mate just once ('no risk'), (2) mate twice but have not yet mated ('future risk'), and (3) mate twice and have already mated ('past risk'). The ESS is to ejaculate minimally with 'no risk' females, and to expand equally with 'past' and 'future' risk females; the direction of the competitive loading is again unimportant. Expenditure again increases with risk, but is now non-zero at extremely low risk. Model 3 examines three cases of partial information where males can identify only one of the three risk states and cannot distinguish between the other two: they therefore have just two information sets or 'contexts'. Expenditure in both contexts typically rises non-linearly from zero with q, but (whatever the loading direction) expenditure is higher in the context with higher risk (e.g. if contexts are 'mated' and 'virgin', males spend more with mated females). However, in highly loaded raffles, sperm expenditure can decrease over part of the range of risk. Also, the direction of the loading now affects expenditure. Biological evidence for the predictions of the models is summarized and discussed.

Matthew J. G. Gage

Papers

Sperm competition, male prudence and sperm-limited females

Sperm competition in fishes: the evolution of testis size and ejaculate characteristics.

Spermatozoal traits and sperm competition in Atlantic salmon: relative sperm velocity is the primary determinant of fertilization success.

Associations between Body Size, Mating Pattern, Testis Size and Sperm Lengths across Butterflies

Sperm competition games: a prospective analysis of risk assessment