Re‐evaluating the distribution of cooperative breeding in birds: is it tightly linked with altriciality?

The evolution of cooperative breeding (CB) in birds has aroused intensive interest for decades, largely due to the paradox that some adults forgo independent breeding to help others. While much effort has been directed at understanding the adaptive significance of CB behavior, much less effort has been spent on understanding its origin. Ligon and Burt argued that the evolution of altriciality played a key role in the origin of CB since CB occurs more frequently in altricial lineages than expected if developmental mode and CB evolved independently and that both traits arose early in the avian tree of life. We mapped presence or absence of CB, and precocial or altricial development on a recent phylogeny of all birds to re‐evaluate their conclusions. Our results suggest altriciality may be more recently derived than previously thought, and that CB species clustered in a derived land bird clade (especially within Passeriformes) where we reconstructed many gains and losses. We did find a link between cooperative breeding and altriciality. However, since CB also occurs in precocial species, has not evolved in many altricial clades, and may have evolved prior to altriciality (based on some classifications of which species have CB), it is not clear whether altriciality is linked to other factors, such as benefits to group living, that are necessary for the acquisition of CB behavior, or whether altriciality may have been a driving force in the evolution of CB itself. The relative importance of these other factors versus altriciality for the origin of CB needs to be considered.     

The breeding system of Ramosmania heterophylla–dioecy or heterostyly?

OWENS, S.J., JACKSON, A., MAUNDER, M., RUDALL, P.JOHNSON, M.A.T., 1993. The breeding system of Ramosmania heterophylla– dioecy or heterostyly?Ramosmania heterophylla (Cafe Marron), endemic to Rodrigues, Mauritius, is now almost extinct, since it appears to exist in the wild as a single plant, but has recently been successfully established at Kew by means of vegetative cuttings. Despite plenty of flowers on the Kew plants, attempts to set seed by manual pollination for seed banking purposes have failed. This investigation showed that pollen is viable and that embryo sacs appear fully developed. Lack of seed set is a result of the non-functional stigmas on short styles. The most likely explanation is that the last-known plant is male, although heterostyly or a mutant stylar developmental gene cannot be ruled out. The plant has a chromosome number of 2n= 22.     

Was monogamy a key step on the hominin road? reevaluating the monogamy hypothesis in the evolution of cooperative breeding

Because human mothers routinely rely on others to help raise their young, humans have been characterized as cooperative breeders.^1–9 Several large‐scale phylogenetic analyses have presented compelling evidence that monogamy preceded the evolution of cooperative breeding in a wide variety of nonhuman animals.^10–14 These studies have suggested that monogamy provides a general rule (the monogamy hypothesis) for explaining evolutionary transitions to cooperative breeding.¹⁵ Given the prevalence of cooperative breeding in contemporary human societies, we evaluate whether this suggests a monogamous hominin past.     

What is cooperative breeding in mammals and birds? Removing definitional barriers for comparative research

Cooperative breeding (i.e. when alloparents care for the offspring of other group members) has been studied for nearly a century. Yet, inconsistent definitions of this breeding system still hamper comparative research. Here, we identify two major inconsistencies, discuss their consequences and propose a way forward. First, some researchers restrict the term ‘cooperative breeding’ to species with non-breeding alloparents. We show that such restrictive definitions lack distinct quantitative criteria to define non-breeding alloparents. This ambiguity, we argue, reflects the reproductive-sharing continuum among cooperatively breeding species. We therefore suggest that cooperative breeding should not be restricted to the few species with extreme reproductive skew and should be defined independent of the reproductive status of alloparents. Second, definitions rarely specify the type, extent and prevalence of alloparental care required to classify species as cooperative breeders. We thus analysed published data to propose qualitative and quantitative criteria for alloparental care. We conclude by proposing the following operational definition: cooperative breeding is a reproductive system where >5% of broods/litters in at least one population receive species-typical parental care and conspecifics provide proactive alloparental care that fulfils >5% of at least one type of the offspring's needs. This operational definition is designed to increase comparability across species and disciplines while allowing to study the intriguing phenomenon of cooperative breeding as a behaviour with multiple dimensions.     

The evolution of photosynthesis…again?

‘Replaying the tape’ is an intriguing ‘would it happen again?’ exercise. With respect to broad evolutionary innovations, such as photosynthesis, the answers are central to our search for life elsewhere. Photosynthesis permits a large planetary biomass on Earth. Specifically, oxygenic photosynthesis has allowed an oxygenated atmosphere and the evolution of large metabolically demanding creatures, including ourselves. There are at least six prerequisites for the evolution of biological carbon fixation: a carbon-based life form; the presence of inorganic carbon; the availability of reductants; the presence of light; a light-harvesting mechanism to convert the light energy into chemical energy; and carboxylating enzymes. All were present on the early Earth. To provide the evolutionary pressure, organic carbon must be a scarce resource in contrast to inorganic carbon. The probability of evolving a carboxylase is approached by creating an inventory of carbon-fixation enzymes and comparing them, leading to the conclusion that carbon fixation in general is basic to life and has arisen multiple times. Certainly, the evolutionary pressure to evolve new pathways for carbon fixation would have been present early in evolution. From knowledge about planetary systems and extraterrestrial chemistry, if organic carbon-based life occurs elsewhere, photosynthesis—although perhaps not oxygenic photosynthesis—would also have evolved.     

The evolution of trade‐offs: where are we?

Trade-offs are a core component of many evolutionary models, particularly those dealing with the evolution of life histories. In the present paper, we identify four topics of key importance for studies of the evolutionary biology of trade-offs. First, we consider the underlying concept of 'constraint'. We conclude that this term is typically used too vaguely and suggest that 'constraint' in the sense of a bias should be clearly distinguished from 'constraint' in the sense of proscribed combinations of traits or evolutionary trajectories. Secondly, we address the utility of the acquisition-allocation model (the 'Y-model'). We find that, whereas this model and its derivatives have provided new insights, a misunderstanding of the pivotal equation has led to incorrect predictions and faulty tests. Thirdly, we ask how trade-offs are expected to evolve under directional selection. A quantitative genetic model predicts that, under weak or short-term selection, the intercept will change but the slope will remain constant. Two empirical tests support this prediction but these are based on comparisons of geographic populations: more direct tests will come from artificial selection experiments. Finally, we discuss what maintains variation in trade-offs noting that at present little attention has been given to this question. We distinguish between phenotypic and genetic variation and suggest that the latter is most in need of explanation. We suggest that four factors deserving investigation are mutation-selection balance, antagonistic pleiotropy, correlational selection and spatio-temporal variation, but as in the other areas of research on trade-offs, empirical generalizations are impeded by lack of data. Although this lack is discouraging, we suggest that it provides a rich ground for further study and the integration of many disciplines, including the emerging field of genomics.     

The evolution of mobile DNAs: when will transposons create phylogenies that look as if there is a master gene?

Some families of mammalian interspersed repetitive DNA, such as the Alu SINE sequence, appear to have evolved by the serial replacement of one active sequence with another, consistent with there being a single source of transposition: the "master gene." Alternative models, in which multiple source sequences are simultaneously active, have been called "transposon models." Transposon models differ in the proportion of elements that are active and in whether inactivation occurs at the moment of transposition or later. Here we examine the predictions of various types of transposon model regarding the patterns of sequence variation expected at an equilibrium between transposition, inactivation, and deletion. Under the master gene model, all bifurcations in the true tree of elements occur in a single lineage. We show that this property will also hold approximately for transposon models in which most elements are inactive and where at least some of the inactivation events occur after transposition. Such tree shapes are therefore not conclusive evidence for a single source of transposition.     

The evolution of the neocortex in mammals: how is phenotypic diversity generated?

Evolution of the mammalian neocortex is difficult to examine directly. For this reason, comparative studies and developmental studies are the best way of gaining insight into the evolutionary process. Comparative studies indicate that neocortical evolution is constrained, and that the types of systems-level modifications made to the neocortex are limited. Developmental studies of gene expression suggest that genetic contingencies set up aspects of cortical organization and connectivity, and that the complex spatial and temporal interactions of genes constrain development and evolution. Although genes obviously contribute to phenotypic variability, variability can also be achieved through alterations in the sensory receptor arrays, or changes in sensory driven activity. The intracellular mechanisms that enable phenotypic variability might evolve, but often the phenotypic characteristic in question is context-dependent.     

When is bigger better? The effects of group size on the evolution of helping behaviours

Understanding the evolution of sociality in humans and other species requires understanding how selection on social behaviour varies with group size. However, the effects of group size are frequently obscured in the theoretical literature, which often makes assumptions that are at odds with empirical findings. In particular, mechanisms are suggested as supporting large‐scale cooperation when they would in fact rapidly become ineffective with increasing group size. Here we review the literature on the evolution of helping behaviours (cooperation and altruism), and frame it using a simple synthetic model that allows us to delineate how the three main components of the selection pressure on helping must vary with increasing group size. The first component is the marginal benefit of helping to group members, which determines both direct fitness benefits to the actor and indirect fitness benefits to recipients. While this is often assumed to be independent of group size, marginal benefits are in practice likely to be maximal at intermediate group sizes for many types of collective action problems, and will eventually become very small in large groups due to the law of decreasing marginal returns. The second component is the response of social partners on the past play of an actor, which underlies conditional behaviour under repeated social interactions. We argue that under realistic conditions on the transmission of information in a population, this response on past play decreases rapidly with increasing group size so that reciprocity alone (whether direct, indirect, or generalised) cannot sustain cooperation in very large groups. The final component is the relatedness between actor and recipient, which, according to the rules of inheritance, again decreases rapidly with increasing group size. These results explain why helping behaviours in very large social groups are limited to cases where the number of reproducing individuals is small, as in social insects, or where there are social institutions that can promote (possibly through sanctioning) large‐scale cooperation, as in human societies. Finally, we discuss how individually devised institutions can foster the transition from small‐scale to large‐scale cooperative groups in human evolution.     

How symbiogenic is evolution?

When new entities are formed by the integration of individual organisms, these new entities possess characteristics which go beyond the sum of the individual properties of each element of the association, resulting in the development of new attributes and capacities as an integrated whole. In this process, these new entities also agglutinate and dynamize synergies not present in the individual organisms. In this sense, evolution is a dynamic process that evolves not in the way of perfection or progress, but in the way of adaptation to new conditions. Symbiogenesis, as an evolutionary mechanism, allows a coherent conceptual rupture with some evolutionary ideas of the past and, at the same time, shows and builds a new approach to life, based on solid evolutionary ideas, expanding evolution to an adequate level of integration with the more recent data in biology. These ideas and concepts should be integrated in a post-neodarwinian approach to evolution that needs further attention from the scientific community. The development of a Symbiogenic Theory of Evolution could contribute toward a new epistemological approach of the symbiotic phenomenon in the evolutionary context. This, in our point of view, could be the beginning of a new paradigm in science that rests almost unexplored.     

Eradication of scrapie with selective breeding: are we nearly there?

Background  

Following EU decision 2003/100/EC Member States have recently implemented sheep breeding programmes to reduce the prevalence of sheep with TSE susceptible prion genotypes. The present paper investigates the progress of the breeding programme in the Netherlands. The PrP genotype frequencies were monitored through time using two sets of random samples: one set covers the years 2005 to 2008 and is taken from national surveillance programme; the other is taken from 168 random sheep farms in 2007. The data reveal that although the level of compliance to the breeding programme has been high, the frequency of susceptible genotypes varies substantially between farms. The 168 sheep farms are a subset of 689 farms participating in a postal survey inquiring about management and breeding strategies. This survey aimed to identify how much these strategies varied between farms, in order to inform assessment of the expected future progress towards eradication of classical scrapie.     

Bias? What bias? The evolution of downstream larval‐feeding in animals

The homology of larval-feeding modes has been a major element in recent discussions about animal phylogeny. 'Downstream-feeding' is one of the two main larval-feeding modes, and is usually equated to an opposed-band system involving ciliary bands called the prototroch and metatroch. Larval-feeding in Spiralia is reviewed here and the homology hypothesis of downstream larval-feeding is expanded, encompassing any feeding involving the prototroch. It is often argued that the presence of planktotrophic larvae using downstream-feeding is plesiomorphic among spiralian animals, and that there is a bias in transformations, such that feeding larvae tend to be lost rather than gained. These hypotheses are assessed in relation to the Spiralia, Trochozoa and particularly polychaete annelids. Cladistic parsimony analyses are performed based on datasets of Rouse & Fauchald (1997) and Rouse (1999) , with an additional character based on downstream larval-feeding. Methods adopted to assess the possibility of a bias in transformations towards loss of downstream larval-feeding include: expanded primary homology arguments, character transformations favouring reversals and polymorphic terminals coded as having downstream larval-feeding. These measures all tend to favour the possibility that downstream larval-feeding will be plesiomorphic, and tends to be lost rather than gained. Nevertheless, all analyses show that downstream larval-feeding is not plesiomorphic, and appears to have evolved multiple times. The results support a conclusion that the prototroch has become associated with feeding a number of times. Hypotheses of metazoan phylogeny that are predicated on the assumption that downstream-feeding larvae are plesiomorphic (e.g. Peterson et al . 1997 ) should be re-assessed.     

The evolution of ADHD: a disorder of communication?

Attention deficit hyperactivity disorder (ADHD) is the most commonly diagnosed psychiatric condition. Many believe that the central disability is impaired inhibition, which leads to reduced abilities in social skills, self-control, organization and time management. The behaviors identified by clinicians as problematic--inattention, hyperactivity and impulsivity--have been incorporated into several evolutionary models as selectively adaptive cognitive skills for surviving the challenges of a variable Pleistocene environment. We propose that the "disabilities" exhibited by individuals with ADHD are maladaptive, and we concur with Barkley that there is a central impairment in the behavioral inhibition system. The underlying neural anatomy and physiology support the possibility that neurotransmitter pathology may have an impact on other interlinked systems (including language), and may also account for the frequent comorbidity of aggression, anxiety, depression, and learning disabilities (many of which are language-related). Language skills compete with other cognitive activities for the attentional system, and thus the evolution of language could not in fact be independent of the evolution of attention. If language represents the ultimate expression of the attentional system, and some individuals with ADHD are seriously impaired in the coordination of interlinked neural systems (including language), then ADHD fits Jerome Wakefield's definition of "harmful dysfunction," and communication impairments should be investigated more thoroughly by clinicians.