Sex chromosomes can differ between species as a result of evolutionary turnover, a process that can be driven by evolution of the sex determination pathway. Canonical models of sex chromosome turnover hypothesize that a new master sex determining gene causes an autosome to become a sex chromosome or an XY chromosome pair to switch to a ZW pair (or vice versa). Here, a novel paradigm for the evolution of sex determination and sex chromosomes is presented, in which there (...) is an evolutionary transition in the master sex determiner, but the X chromosome remains unchanged. There are three documented examples of the novel paradigm, and it is hypothesized that a similar process could happen in a ZW sex chromosome system. Three other taxa are also identified where the novel paradigm may have occurred, and how it could be distinguished from canonical trajectories in these and additional taxa is also described. (shrink)

The evolution of sex in eukaryotes represents a paradox, given the “twofold” fitness cost it incurs. We hypothesize that the mutational dynamics of the mitochondrial genome would have favored the evolution of sexual reproduction. Mitochondrial DNA (mtDNA) exhibits a high‐mutation rate across most eukaryote taxa, and several lines of evidence suggest that this high rate is an ancestral character. This seems inexplicable given that mtDNA‐encoded genes underlie the expression of life's most salient functions, including energy conversion. We propose (...) that negative metabolic effects linked to mitochondrial mutation accumulation would have invoked selection for sexual recombination between divergent host nuclear genomes in early eukaryote lineages. This would provide a mechanism by which recombinant host genotypes could be rapidly shuffled and screened for the presence of compensatory modifiers that offset mtDNA‐induced harm. Under this hypothesis, recombination provides the genetic variation necessary for compensatory nuclear coadaptation to keep pace with mitochondrial mutation accumulation. (shrink)

The evolution of sexual reproduction is a striking case of explanatory pluralism, meaning that one needs to refer to more than one explanation in order to adequately account for it. I develop the concept a domain of phenomena in order to analysis this pluralism. Pluralism exists when a phenomenon can be included in more that one homogeneous domain or in a heterogeneous domain. I argue that in some cases domain partitioning can be used to decrease pluralism, but that in (...) the case of sex domains are overlapping and interconnecting, or in other words bear an orthogonal relationship to one another, and hence cannot be partitioned in such a way as to eliminate pluralism. (shrink)

Few topics have intrigued biologists as much as the evolution of sex. Understanding why sex persists despite its costs requires not just rigorous theoretical study, but also empirical data on related fundamental issues, including the nature of genetic variance for fitness, patterns of genetic interactions, and the dynamics of adaptation. The increasing feasibility of examining genomes in an experimental context is now shedding new light on these problems. Using this approach, McDonald et al. recently demonstrated that sex uncouples beneficial (...) and deleterious mutations, allowing selection to proceed more effectively with sex than without. Here we discuss the insights provided by this study, along with other recent empirical work, in the context of the major theoretical models for the evolution of sex. (shrink)

Sex chromosomes in plants have been known for a century, but only recently have we begun to understand the mechanisms behind sex determination in dioecious plants. Here, we discuss evolution of sex determination, focusing on Silene latifolia, where evolution of separate sexes is consistent with the classic “two mutations” model—a loss of function male sterility mutation and a gain of function gynoecium suppression mutation, which turned an ancestral hermaphroditic population into separate males and females. Interestingly, the gynoecium suppression (...) function in S. latifolia evolved via loss of function in at least two sex‐linked genes and works via gene dosage balance between sex‐linked, and autosomal genes. This system resembles X/A‐ratio‐based sex determination systems in Drosophila and Rumex, and could represent a steppingstone in the evolution of X/A‐ratio‐based sex determination from an active Y system. (shrink)

Why do the two sexes have different lifespans and rates of aging? Two hypotheses based on asymmetric inheritance of sex chromosomes (“unguarded X”) or mitochondrial genomes (“mother's curse”) explain sex differences in lifespan as sex‐specific maladaptation leading to increased mortality in the shorter‐lived sex. While asymmetric inheritance hypotheses equate long life with high fitness, considerable empirical evidence suggests that sexes resolve the fundamental tradeoff between reproduction and survival differently resulting in sex‐specific optima for lifespan. However, selection for sex‐specific values in (...) life‐history traits is constrained by intersexual genetic correlations resulting in intra‐locus sexual conflict over optimal lifespan. The available data suggest that the evolution of sexual dimorphism only partially resolves these conflicts. Sexual conflict over optimal trait values, which has been demonstrated in model organisms and in humans, is likely to play a key role in shaping the evolution of lifespan, as well as in maintaining genetic variation for sex‐specific diseases. (shrink)

Sex is determined by non‐Mendelian genetic elements overriding the sex factors carried by the heterochromosomes in some species of terrestrial isopods. A bacterium Wolbachia and a non‐bacterial feminizing factor (f) can both force chromosomal males of Armadillidium vulgare to become phenotypic functional females. The f factor is believed to be a genetic element derived from the Wolbachia genome that becomes inserted into the host nuclear genome. The feminizing factors can be considered to be selfish genetic elements because they bias their (...) host's sex ratio to increase their own transmission. New sex‐determining genes are selected (genes resisting the feminizing effects, or the transmission of feminizing elements) as a consequence of the conflict between these elements and the rest of the host's genome. These events drive the sex‐determining mechanisms to evolve, and may explain the polymorphism of sex factors and the poor differentiation of the heterochromosomes in isopods. (shrink)

Frequent independent origins of environmental sex determination (ESD) are assumed within amniotes. However, the phylogenetic distribution of sex‐determining modes suggests that ESD is likely very ancient and may be homologous across ESD groups. Sex chromosomes are demonstrated to be old and stable in endothermic (mammals and birds) and many ectothermic (non‐avian reptiles) lineages, but they are mostly non‐homologous between individual amniote lineages. The phylogenetic pattern may be explained by ancestral ESD with multiple transitions to later evolutionary stable genotypic sex determination. (...) It is pointed out here that amniote ESD shares several key aspects with sequential hermaphroditism of fishes such as a lack of sex differences in genomes, biased population sex ratios, and potentially also molecular mechanism related to general stress responses. Here, it is speculated that ESD evolves via a heterochronic shift of the sensitive period of sex change from the adult to the embryonic stage in a hermaphroditic amniote ancestor. Also see the video abstract here https://youtu.be/q2mjtlCefu4. (shrink)

Benenson et al. provide a compelling case for treating greater investment into self-protection among females as an adaptive strategy. Here, we wish to expand their proposed adaptive explanation by placing it squarely in modern state-based and behavioural life-history theory, drawing on Veit'spathological complexityframework. This allows us to make sense of alternative “lifestyle” strategies, rather than pathologizing them.

The evolutionary maintenance of sexual reproduction is a case of explanatory pluralism of central importance to evolutionary biology. I analyze this pluralism from an epistemological perspective. My thesis is that the various explanations of sex are explanatory by virtue of local factors and hence are importantly distinct from one another and cannot be subsumed under a single unifying framework. A critic may argue that philosophical accounts of mechanism can provide just such a framework. I show that this attempt at unification (...) fails because the accounts of sex are not explanatory simply in light of being their being mechanistic; rather they are explanatory because they are particular kinds of mechanisms. The explanatory power of a mechanism is dependent on its epistemological context and the contexts of these mechanisms differ. (shrink)

In most discussions of the evolution of sex chromosomes, it is presumed that the morphological differences between the X and Y were initiated by genetic changes. An alternative possibility is that, in the early stages, a key role was played by epigenetic modifications of chromatin structure that did not depend directly on genetic changes. Such modifications could have resulted from spontaneous epimutations at a sex‐determining locus or, in mammals, from selection in females for the epigenetic silencing of imprinted regions (...) of the paternally derived sex chromosome. Other features of mammalian sex chromosomes that are easier to explain if the epigenetic dimension of chromosome evolution is considered include the relatively large number of X‐linked genes associated with human brain development, and the overrepresentation of spermatogenesis genes on the X. Both may be evolutionary consequences of dosage compensation through X‐inactivation. BioEssays 26:1327–1332, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

Sexual reproduction is the dominant reproductive mode in eukaryotes but, in many taxa, it has never been observed. Molecular methods that detect evidence of sex are largely based on the genetic consequences of sexual reproduction. Here we describe a powerful new approach to directly search genomes for genes that function in meiosis. We describe a “meiosis detection toolkit”, a set of meiotic genes that represent the best markers for the presence of meiosis. These genes are widely present in eukaryotes, function (...) only in meiosis and can be isolated by degenerate PCR. The presence of most, or all, of these genes in a genome would suggest they have been maintained for meiosis and, implicitly, sexual reproduction. In contrast, their absence would be consistent with the loss of meiosis and asexuality. This approach will help to understand both meiotic gene evolution and the capacity for meiosis and sex in putative obligate asexuals. BioEssays 30:579–589, 2008. © 2008 Wiley Periodicals, Inc. (shrink)

Various explanations have been proposed to account for complex differentiation and development in humans, despite the human genome containing only two to three times the number of genes in invertebrates. Ignored are the actions of adrenal and sex steroids—androgens, estrogens, glucocorticoids, mineralocorticoids, and progestins—which act through receptors that arose from an ancestral nuclear receptor in a protochordate. This ligand‐based mechanism is unique to vertebrates and was integrated into the already robust network of transcription factors in invertebrates. Adrenal and sex steroids (...) influence almost all aspects of vertebrate differentiation and development. I propose that evolution of this ligand‐based mechanism in a primitive vertebrate was an important contribution to vertebrate complexity. Sequencing of genomes from a cephalochordate, such as amphioxus, and from hagfish and lamprey will establish early events in the evolution of steroid hormone signaling, and also allow genetic studies to elucidate how vertebrate complexity depends on steroid hormones. BioEssays 25:396–400, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

Sexual selection traditionally involves male-male competition and female choice, but in some species, including humans, sexual selection can also involve female-female competition and male choice. The degree to which one aspect of sexual selection or another is manifest in human populations will be influenced by a host of social and ecological variables, including the operational sex ratio. These variables are discussed in connection with the relative contribution of sexual selection and the division of labor to the evolution of human (...) sex differences. (shrink)

Sex, Gender, Ethics, and the Darwinian Evolution of Humanity examines the impact of Darwin's 'Descent of Man' on contemporary biology and the humanities. Its publication in 1871 was a founding event in anthropology. Its content was primarily concerned with the development of sexual life, social life, and intellectual life, not only as outcomes of evolution, but as components that have actively intermixed over time with the evolutionary mechanism of natural selection. The stamp of Darwinism on modern thought is (...) still very important and brings novelties to academic studies. Several fields influenced by Darwinian anthropology developed in recent decades, including evolutionary ethics, the evolution of sociality and sexual communication in animal and plant species. Sociobiology and evolutionary psychology are topics that draw heavily on Darwin's 'Descent of Man'. The understanding of Darwin's thought has also progressed greatly in recent decades, following the systematic study of Darwin's correspondence and notebooks, leading to a reassessment of the development of his thought on humans, social groups, and heredity, and how they come together in his theory of evolution. The book combines a historical perspective on Darwin's achievement and his legacy. It will be of interest to students and scholars in a variety of fields, from experimental biology to the social and historical sciences. (shrink)

theories. What was Freud thinking, when, and why and what were the major influences which shaped his ideas? We follow the inner movement of his theory construction, its meaning and coherence, as well as his conceptual logic and personal directions concerning his evolving views of the reciprocal interactions between mind and body, the motivational force of instinctual drives, and the dominant role of sexuality rooted in evolutionary biology in human development, behaviour, and the creation of neurotic disturbances. We follow Freud's (...) construction and sequential reconstructions of his theoretical models concerning the nature, dynamics, and principles of unconscious mental functioning, including his changing concepts on the nature and purpose of dreams. We trace his changing views on the role of deferred action of early childhood experiences and the determining role of unconscious fantasy, psychic reality, in the formation of adult character structure and neuroses. Through such historical analysis this book provides grounding for a meaningful understanding of Freud's familiar concepts: id, ego, superego, and the Oedipus complex. We explore what these concepts meant to Freud, why he conceived them, and what functions they served in his theory of mind. This is the perfect book for students and trainees wanting to learn more about the development of Freud's ideas, as well as for established psychoanalysts and psychotherapists interested in expanding their knowledge of Freud's theories. (shrink)

Brian Skyrms has studied the evolutionary dynamics of a simple bargaining game. Fair-dealing is the strategy 'demand 1/2', competing with the more modest strategy 'demand 1/3' and the greedier strategy 'demand 2/3'. Individuals leave offspring in proportion to their accumulated payoffs. The rules for payoffs from encounters penalize low- and high-demanders. The result is a significant basin of attraction for fair-dealing as an evolutionarily stable strategy. From these considerations Skyrms concludes that a disposition to fair-dealing could have evolved. A very (...) different picture emerges, however, when one considers genetic bases for the dispositions involved. A simple two-allele sexual model produces very different stable equilibria in the distribution of behavioral phenotypes. The equilibria for Skyrms's purely phenotypic selection process will not in general be attainable once one enters some simple genetic considerations. (shrink)

A hypothesis on the evolutionary origin of the genetic pathway of sex determination in the nematode Caenorhabditis elegans is presented here. It is suggested that the pathway arose in steps, driven by frequency‐dependent selection for the minority sex at each step, and involving the sequential acquisition of dominant negative, neomorphic genetic switches, each one reversing the action of the previous one. A central implication is that the genetic pathway evolved in reverse order from the final step in the hierarchy up (...) to the first. The possible applicability of the model to the other well‐characterized sex determination pathway, that of Drosophila melanogaster, and to sex determination in mammals, is discussed, along with some potential implications for pathway evolution in general. Finally, the specific molecular and population genetic questions that the model raises are described and some tests are proposed. (shrink)

Gradual degradation seems inevitable for non‐recombining sex chromosomes. This has been supported by the observation of degenerated non‐recombining sex chromosomes in a variety of species. The human Y chromosome has also degenerated significantly during its evolution, and theories have been advanced that the Y chromosome could disappear within the next ∼5 million years, if the degeneration rate it has experienced continues. However, recent studies suggest that this is unlikely. Conservative evolutionary forces such as strong purifying selection and intrachromosomal repair (...) through gene conversion balance the degeneration tendency of the Y chromosome and maintain its integrity after an initial period of faster degeneration. We discuss the evidence both for and against the extinction of the Y chromosome. We also discuss potential insights gained on the evolution of sex‐determining chromosomes by studying simpler sex‐determining chromosomal regions of unicellular and multicellular microorganisms. (shrink)

Although I do not know Richard Doyle personally, I would say that Darwin’s Pharmacy: Sex Plants and the Evolution of the Noosphere is a deeply personal book. Not only does the author offer multiple accounts of his own multicontinental explorations of intraspecies cross-pollination, but he also provides many rhetorical analyses of trip reports, biological treatises, and science fiction, all of which seem to be crucial constitutive elements of his research. That is, this is not a book that offers abstract (...) erudition—though there is plenty of content that anyone can extract from it—but one that offers something more rare. Here, I am reminded of Goethe’s famous remark, which Nietzsche chose to use as the epigraph to.. (shrink)

The question of what role sex plays in evolution is still open despite decades of research. It has often been assumed that sex should facilitate the increase in fitness. Hence, the fact that it may break down highly favorable genetic combinations has been seen as a problem. Here, we consider an alternative approach. We define a measure that represents the ability of alleles to perform well across different combinations and, using numerical iterations within a classical population-genetic framework, show that (...) selection in the presence of sex favors this ability in a highly robust manner. We also show that the mechanism responsible for this effect has been out of the purview of previous theory, because it operates during the evolutionary transient, and that the breaking down of favorable genetic combinations is an integral part of it. Implications of these results and more to evolutionary theory are discussed. (shrink)

Biologists have long theorized about the evolution of life cycles, meiosis, and sexual reproduction. We revisit these topics and propose that the fundamental difference between life cycles is where and when multicellularity is expressed. We develop a scenario to explain the evolutionary transition from the life cycle of a unicellular organism to one in which multicellularity is expressed in either the haploid or diploid phase, or both. We propose further that meiosis might have evolved as a mechanism to correct (...) for spontaneous whole‐genome duplication (auto‐polyploidy) and thus before the evolution of sexual reproduction sensu stricto (i.e. the formation of a diploid zygote via the fusion of haploid gametes) in the major eukaryotic clades. In addition, we propose, as others have, that sexual reproduction, which predominates in all eukaryotic clades, has many different advantages among which is that it produces variability among offspring and thus reduces sibling competition. (shrink)

The endangered species Tokudaia osimensis has the unique chromosome constitution of 2n = 25, with an XO/XO sex chromosome configuration (2n = 25; XO). There is urgency to preserve this species and to elucidate the regulator(s) that can discriminate the males and females arising from the indistinguishable sex chromosome constitution. However, it is not realistic to examine this rare animal species by sacrificing individuals. Recently, true naïve induced pluripotent stem cells were successfully generated from a female T. osimensis, and the (...) sexual plasticity of its germ cells was elucidated. This achievement constitutes the basis of an attractive research area, including embryonic fate determination, sex determination, and factor(s) that can replace the Y chromosome. In this essay, concrete strategies to conserve rare animal species and to reveal their specific characteristics using other compatible and abundant animals are proposed. -/- . (shrink)

Boehm's model conceptualizes a common ancestor to humans, chimpanzees, and bonobos at several million years B.P., followed by a model of prehistoric foragers at 25,000-50,000 B.P. based on ethnographic data from twentieth-century hunters and gatherers. By putting processes of complex communication into the picture, we can refine Boehm's model considerably by filling in significant scenarios for humans beginning at perhaps 2 million years ago. These include a suite of features that include constraints on sexual behaviour, a rudimentary division of labour, (...) the gendering of morality, socialized control over junior males, increased group size, increased home range size, the reduction of bullying behaviour, and a reverse dominance hierarchy that promotes egalitarianism among fully adult males. Boehm's model emphasizes the last two of these, but it is consistent with the others as well. Cultural or proto-cultural elaboration of beliefs and associated practices is the symbolic dimension of morality and is an important component of human evolution. Boehm's model is extremely important for outlining socio-political mechanisms and concomitants in the formation of adult male egalitarian groups, and, in the process, identifying a key element in the evolution of human morality. (shrink)

The evolution of nonconceptive sexuality in bonobos and chimpanzees is discussed from a functional perspective. Bonobos and chimpanzees have three functions of sexual activity in common (paternity confusion, practice sex, and exchange for favors), but only bonobos use sex purely for communication about social relationships. Bonobo hypersexuality appears closely linked to the evolution of female-female alliances. I suggest that these alliances were made possible by relaxed feeding competition, that they were favored through their effect on reducing sexual coercion, (...) and that they are ultimately responsible for the relaxed social conditions that allowed the evolution of “communication sex.”. (shrink)

Despite a revived interest in explaining the evolution of anisogamy in recent years (i.e. different—micro and macrogametes), there remain more questions than answers. The topic is important because it is thought to be the foundation of the theory of gender differences and relations. Twelve of these questions are briefly reviewed here—(1) the distinction between sex and sexual types; (2) the distinction between mating types and anisogamy; (3) the possible role of ecological as well as social evolution in proto-gender (...) differences and relations; (4) the life history strategies involved; (5) whether the relevant social relationships are based on conflict or cooperation; (6) the origin and properties of meiosis; (7) the conformity of theories with sex allocation theory; (8) the relevance of multiple levels of selection; (9) appropriate modelling strategies; (10) the relationship between the kind of gametes produced and secondary sexual characteristics; (11) the relevance to humans; and (12) how realistic the search for a single explanation is. It is concluded that polarized sexuality may be a form of trade. (shrink)

During human evolutionary history, there were “trade-offs” between expending time and energy on child-rearing and mating, so both men and women evolved conditional mating strategies guided by cues signaling the circumstances. Many short-term matings might be successful for some men; others might try to find and keep a single mate, investing their effort in rearing her offspring. Recent evidence suggests that men with features signaling genetic benefits to offspring should be preferred by women as short-term mates, but there are trade-offs (...) between a mate's genetic fitness and his willingness to help in child-rearing. It is these circumstances and the cues that signal them that underlie the variation in short- and long-term mating strategies between and within the sexes. Key Words: conditional strategies; evolutionary psychology; fluctuating asymmetry; mating; reproductive strategies; sexual selection. (shrink)

This book inquires into the swarm of ontological, epistemological, and ethical questions provoked by psychedelic experience in the context of global ecological crisis. Richard M. Doyle is professor of English and science, technology, and society at Pennsylvania State University. He is the author of On Beyond Living and Wetwares.

Patterns in the data on human sexuality support the hypothesis that the bases of sexual emotions are products of natural selection. Most generally, the universal existence of laws, rules, and gossip about sex, the pervasive interest in other people's sex lives, the widespread seeking of privacy for sexual intercourse, and the secrecy that normally permeates sexual conduct imply a history of reproductive competition. More specifically, the typical differences between men and women in sexual feelings can be explained most parsimoniously as (...) resulting from the extraordinarily different reproductive opportunities and constraints males and females normally encountered during the course of evolutionary history. Men are more likely than women to desire multiple mates; to desire a variety of sexual partners; to experience sexual jealousy of a spouse irrespective of specific circumstances; to be sexually aroused by the sight of a member of the other sex; to experience an autonomous desire for sexual intercourse; and to evaluate sexual desirability primarily on the bases of physical appearance and youth.The evolutionary causes of human sexuality have been obscured by attempts to find harmony in natural creative processes and human social life and to view sex differences as complementary. The human female's capacity for orgasm and the loss of estrus, for example, have been persistently interpreted as marriage-maintaining adaptations. Available evidence is more consistent with the view that few sex differences in sexuality are complementary, that many aspects of sexuality undermine marriage, and that sexuality is less a unifying than a divisive force in human affairs. (shrink)