Gyrencephaly (the folding of the surface of the neocortex) is a mammalian-specific trait present in almost all mammalian orders. Despite the widespread appearance of the trait, little is known about the mechanism of its genesis or its adaptive significance. Still, most of the hypotheses proposed concentrated on the pattern of connectivity of mature neurons as main components of gyri formation. Recent work on embryonic neurogenesis in several species of mammals revealed different progenitor and stem cells and their neurogenic potential as having important roles in the process of gyrification. Studies in the field of comparative neurogenesis revealed that gyrencephaly is an evolutionarily labile trait, and that some species underwent a secondary loss of a convoluted brain surface and thus reverted to a more ancient form, a less folded brain surface (lissencephaly). This phenotypic reversion provides an excellent system for understanding the phenomenon of secondary loss. In this review, we will outline the theory behind secondary loss and, as specific examples, present species that have undergone this transition with respect to neocortical folding. We will also discuss different possible pathways for obtaining (or losing) gyri. Finally, we will explore the potential adaptive consequence of gyrencephaly relative to lissencephaly and vice versa.

Les asimetries funcionals manuals han estat àmpliament estudiades en els primats no humans i altres espècies animals durant les últimes dues dècades. Resulta especialment interessant observar si l'especialització hemisfèrica cerebral, tan característica de l'ésser humà, està o no present i en quin grau dins del regne animal. En ximpanzés, trobem pocs treballs en què s'hagin avaluat les preferències manuals en una única mostra des d'un punt de vista observacional i experimental conjuntament, i cap en ambients intermedis (naturalitzats). El nostre objectiu ha estat comparar els patrons de preferència manual en situacions espontànies i experimentals en una mateixa mostra de ximpanzés i veure quines situacions incideixen sobre la direcció, intensitat i grau d'asimetria individual versus poblacional en els individus. En la fase observacional es van estudiar 53 comportaments espontanis unimanuals i bimanuals que feien referència a contextos tròfics, manipulatius-exploratoris, autodirigits, posturals i socials, entre d'altres. En la fase experimental es van avaluar dues tasques: una unimanual (simple reaching: recollida d'ítems petits d'aliment amb prensions de precisió) i una altra bimanual (tube task: extracció bimanual d'aliment d'un tub). Durant la fase observacional es van dur a terme 2.226 sessions (2005 a 2008: 33 mesos). La fase experimental va tenir una durada total de 4 mesos (2007). La mostra estava formada per un grup de 14 ximpanzés (Pan troglodytes) allotjats al Centre de Recuperació de Primats de la Fundació Mona (Girona). En ambdues fases es va utilitzar un mostreig focal amb registre activat per transicions. Globalment, tant durant la fase observacional com experimental, la major part de la mostra va presentar preferències individuals laterals de manera significativa. Es van comparar els índexs de lateralitat per a les quatre condicions d'estudi: unimanual-espontani (UES), bimanual-espontani (BES), unimanual-experimental (UEX) i bimanual-experimental (BEX). No es van trobar diferències en la direcció de les preferències entre les quatre condicions tot i que sí en la intensitat de la lateralització. D'aquesta manera, la intensitat de la preferència manual era major en les accions bimanuals que en les unimanuals, i també en les tasques experimentals que en les espontànies. L'ús d'instruments i el tipus de prensió també van augmentar la intensitat de les preferències encara que no van tenir cap efecte sobre la direcció d'aquestes. Es va detectar una asimetria dretana poblacional en les accions bimanuals espontànies però no en les experimentals. També es va detectar una asimetria dreta a nivell poblacional en tenir en compte totes les tasques conjuntament. D'aquesta manera, ha estat la primera ocasió que s'evidencia que els ximpanzés allotjats en ambients intermedis són dretans a nivell poblacional. En conclusió, els ximpanzés presentarien un patró de lateralització homòleg al dels éssers humans, sobretot en aquelles accions que impliquen una complexitat en la seva execució (bimanualitat, instrumentalitat i precisió) i que són les més rellevants per entendre el procés d'hominització conductual d'aquest tret a Homo sapiens. / Las asimetrías funcionales manuales han sido ampliamente estudiadas en los primates no humanos y otras especies animales durante las últimas dos décadas. Resulta especialmente interesante observar si la especialización hemisférica cerebral, tan característica del ser humano, está o no presente y en qué grado dentro del reino animal. En chimpancés, encontramos pocos trabajos en los que se hayan evaluado las preferencias manuales en una única muestra desde un punto de vista observacional y experimental conjuntamente, y ninguno en ambientes intermedios (naturalizados). Nuestro objetivo ha sido comparar los patrones de preferencia manual en situaciones espontáneas y experimentales en una misma muestra de chimpancés y ver qué situaciones inciden sobre la dirección, intensidad y grado de asimetría individual versus poblacional en los individuos. En la fase observacional se estudiaron 53 comportamientos espontáneos unimanuales y bimanuales que hacían referencia a contextos tróficos, manipulativos-exploratorios, autodirigidos, posturales y sociales, entre otros. En la fase experimental evaluaron dos tareas: una unimanual (simple reaching: recogida de ítems pequeños de alimento con prensiones de precisión) y otra bimanual (tube task: extracción bimanual de alimento de un tubo). Durante la fase observacional se llevaron a cabo 2226 sesiones (2005 a 2008: 33 meses). La fase experimental tuvo una duración total de 4 meses (2007). La muestra estaba formada por un grupo de 14 chimpancés (Pan troglodytes) alojados en el Centro de Recuperación de Primates de Fundación Mona (Girona). En ambas fases se utilizó un muestreo focal con registro activado por transiciones. Globalmente, tanto durante la fase observacional como experimental, la mayor parte de la muestra presentó preferencias individuales laterales de manera significativa. Se compararon los índices de lateralidad para las cuatro condiciones de estudio: unimanual-espontáneo (UES), bimanual-espontáneo (BES), unimanual-experimental (UEX) y bimanual-experimental (BEX). No se encontraron diferencias en la dirección de las preferencias entre las cuatro condiciones aunque sí en la intensidad de la lateralización. De esta manera, la intensidad de la preferencia manual era mayor en las acciones bimanuales que en las unimanuales, y también en las tareas experimentales que en las espontáneas. El uso de instrumentos y el tipo de prensión también aumentaron la intensidad de las preferencias aunque no tuvieron ningún efecto sobre la dirección de éstas. Se detectó una asimetría diestra poblacional en las acciones bimanuales espontáneas aunque no en las experimentales. También se detectó una asimetría diestra a nivel poblacional al tener en cuenta todas las tareas conjuntamente. De esta manera, ha sido la primera ocasión que se evidencia que los chimpancés alojados en ambientes intermedios son diestros a nivel poblacional. En conclusión, los chimpancés exhibirían un patrón de lateralización homólogo al de los seres humanos, sobre todo en aquellas acciones que implican una complejidad en su ejecución (bimanualidad, instrumentalidad y precisión) y que son las más relevantes para entender el proceso de hominización conductual de este rasgo en Homo sapiens. / Manual functional asymmetries have been extensively studied in nonhuman primates and other animals for the last two decades. It is particularly interesting to see if brain hemispheric specialization, so characteristic human beings, is present or not and to what extent within the animal kingdom. In chimpanzees, there are few studies that have assessed hand preferences in a single sample from an experimental and observational standpoint, and none have been done within intermediate (naturalized) environments. Our objective was to compare the patterns of hand preference for spontaneous and experimental actions in the same sample of chimpanzees and observe what situations impact on the direction, strength and the degree of individual versus population asymmetry in individuals. In the observational phase, we studied 53 spontaneous unimanual and bimanual behaviors related to feeding contexts, manipulative-exploratory, self-directed, postural and social, among others. In the experimental phase, we evaluated two tasks: unimanual (simple reaching: collection of small items of food with precision grips) and bimanual (tube task: bimanual removal of food from a tube). During the observational phase, we conducted 2.226 sessions (from 2005 to 2008: 33 months). The experimental phase lasted a total of four months (2007). The sample was composed of a group of 14 chimpanzees (Pan troglodytes) housed at the Mona Foundation Primate Rehabilitation Center (Girona). In both phases, a focal sampling method was used. Overall, during both observational and experimental phases, the majority of the sample showed significant individual lateral preferences. We compared the handedness index for four study conditions: unimanual-spontaneous (UES), bimanual-spontaneous (BES), unimanual-experimental (UEX) and bimanual-experimental (BEX). There were no differences in the direction of preferences among the four conditions, but there were differences in the strength of lateralization. The strength of hand preference was higher in bimanual than in unimanual ones, and also in the experimental tasks than in the spontaneous ones. The use of tools and the type of grip also increased the strength of preferences, but had no effect on the direction of such preferences. We detected right-handedness at population-level for spontaneous bimanual actions, but not for the experimental ones. There was also a right asymmetry at the population level when considering all the tasks together. In this way, this was the first evidence that chimpanzees housed in an intermediate environment demonstrate right-handedness at population-level. In conclusion, chimpanzees exhibit a pattern of lateralization homologous to human beings, especially those actions that involve a complexity of execution (bimanual, tool use and precision grip) and that are most relevant for understanding the behavioral process of hominization of this trait in Homo sapiens.

Competition over access for mates has led to the evolution of many striking examples of morphological traits and behaviour in animals. The rapid development of the sexual selection field in recent decades have dramatically advanced our understanding of what traits make individuals more successful in attracting mates and how preferences for mates evolve over time. However, till now, research in this field has put less emphasis on the mechanisms that underlie variation in mate choice and sexual behaviour. Cognitive processes could potentially be key drivers of individual variation in mating preferences and sexual behaviours and therefore deserve further investigation. In this thesis, I used guppies artificially selected for relative brain size as the model system to study the association between brain size, cognitive ability and various aspects of mate choice. Previous studies in this model system showed that large-brained individuals of both sexes outperformed small-brained individuals in cognitive tests. Here I quantified their sexual behaviours and mating preferences to provide novel empirical data concerning the association between brain size, cognitive ability and sexual selection. In dichotomous choice preference tests based on visual cues, comparisons between large-brained and small-brained guppies showed important differences in their assessment of mate quality. These results are not driven by pre-existing visual biases caused by the artificial selection since further investigation of the visual capacity of these fish detected no differences between large-brained and small-brained individuals in their sensitivity to colour or in their capacity to resolve spatial detail. I also quantified sexual behaviour in male guppies artificially selected for relative brain size and found no difference in the behaviours of large-brained and small-brained males in a single male-single female non-competitive scenario. On the contrary, in a more complex social setting I found a reduction in large-brained males in the rate of courtship towards females and dominance displays towards other males when exposed to different degrees of predation threat and different numbers of male competitors. However, this reduction in behavioural intensity did not result in a lower access to copulation with females for large-brained males. I likewise evaluated female sexual behaviour and found that large-brained females had higher behavioural flexibility such that they decreased their receptiveness towards males more strongly under higher levels of predation threat. Together, these results provide novel empirical evidence that brain size and cognitive ability are tightly linked to mating preferences and sexual behaviours. These findings suggest that brain size and cognitive ability might be important mechanisms behind variation in mating preferences and in sexually selected traits across and within species. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 5: Manuscript.</p>