Asymmetric Broca's area in great apes.

Brodmann's area 44 delineates part of Broca's area within the inferior frontal gyrus of the human brain and is a critical region for speech production, being larger in the left hemisphere than in the right - an asymmetry that has been correlated with language dominance. Here we show that there is a similar asymmetry in this area, also with left-hemisphere dominance, in three great ape species (Pan troglodytes, Pan paniscus and Gorilla gorilla). Our findings suggest that the neuroanatomical substrates for left-hemisphere dominance in speech production were evident at least five million years ago and are not unique to hominid evolution.

Interaction between lateralized systems: exploring the complexity of laterality.

To assess the extent of interaction between lateral biases in response systems at different levels of the neuraxis and detect the possible presence of different patterns of interaction related to population subgroups, we investigated laterality in hand reaching, whole-body turning and eye use in 20 bushbabies (Otolemur garnettii). Two subgroups were clearly identified: the STABLE group was composed of subjects, mainly females, that were consistent in hand preference and had correlation of hand/eye bias; the UNSTABLE group included subjects, mainly males, that showed instability in hand preference as a function of change in test conditions and had correlation of hand/turning bias. Results are interpreted to support the value of the study of interaction between lateral biases as a way of gaining a deeper understanding of the complexity of laterality.

Population lateralisation and social behaviour: a study with 16 species of fish.

We investigated turning responses in 16 species of fish faced with a vertical-bar barrier through which a learned dummy predator was visible. Ten of these species showed a consistent lateral bias to turn preferentially to the right or to the left. Species belonging to the same family showed similar directions of lateral biases. We performed an independent test of shoaling tendency and found that all gregarious species showed population lateralisation, whereas only 40% of the non-gregarious species did so. The results provide some support to the Rogers (1989) hypothesis that population lateralisation might have been developed in relation to the need to maintain coordination among individuals in behaviours associated with social life.

Pawedness and motor asymmetries in toads.

Pawedness was investigated in three species of toads, Bufo bufo , Bufo viridis , and Bufo marinus. Samples from natural populations were collected in two successive years and tested during attempts to remove a strip of paper stuck onto the snout ( Bufo bufo and Bufo viridis ) or during attempts to remove an elastic balloon wrapped around the head ( Bufo bufo ). A preferential right forelimb use at the population level was observed in Bufo bufo in both tests, whereas no clear pawedness was observed in Bufo viridis. Bufo marinus also showed no laterality in the paper-strip test, but it appeared to be strongly lateralised in another test that investigated which side the toad rotated when turned on its back underwater. Preferentially turning to the toad's left side as part of a righting response, Bufo marinus released the left forelimb first, using the right forelimb to control the roll to the upright position, and push to the surface. These results suggest that preferential limb use has a long evolutionary history, which dates back to early tetrapods.

Origins and functions of laterality: interactions of motoric systems.

To evaluate lateral motoric bias in response systems at different levels of the neuraxis and assess the extent of interaction between these levels in the small-eared bushbaby (Otolemur garnettii), 27 animals were tested for lateral bias in hand use and whole-body turn bias in two postural conditions. Subjects retrieved mealworms quadrupedally by reaching downwards into glass jars and bipedally by reaching upwards to baited straws. Eye bias was assessed separately. Behaviours were scored from videotape. Two subgroups were identified: SHIFTERS changed hand preference with posture and had correlations of hand/eye bias in quadrupedal posture and of hand/turn bias, with more bimanual reaching, in bipedal posture; NONSHIFTERS were consistent in hand preference and more strongly lateralised in reach and turn than SHIFTERS. Subgroups did not differ in reach efficiency. Results are interpreted to support the value of the analysis of motoric levels and their interactions in the study of the evolution of laterality. Assuming natural selection for coordinated and targeted behaviours to be the source of lateralisation, several proposals in support of a motoric theory of laterality origins and functions are advanced.

Lateral asymmetries during escape behavior in a species of teleost fish (Jenynsia lineata).

Lateral asymmetries in the direction of turning during escape behavior in a species of teleost fish, Jenynsia lineata, are reported. When faced with the visual image of a simulated predator, approximately half of the individuals exhibited a significant bias to turn rightwards or leftwards, and the asymmetry tended to be retained when the same fish were retested 1 month later. Some morphological characters (pectoral fin rays, scales in natural row, supraorbital, preopercular, and postotic pores) were measured to check whether the degree of behavioral asymmetry was correlated with morphological fluctuating asymmetries associated with environmental stress or reduced heterozygosis. The results showed that it was not. The implications of these results for the interpretation of behavioral lateralization at the individual and population level are discussed.

Lateralization of displays during aggressive and courtship behaviour in the Siamese fighting fish (Betta splendens).

Evidence for right-left asymmetries in eye use at the individual level in the Siamese fighting fish, Betta splendens, is reported. When faced with their mirror image (in two daily trials of 10 min each), adult male Betta splendens showed consistency in their right or left eye use during threat lateral displays. Moreover, if one side was preferred by an individual to exhibit the lateral displays, then the duration of the displays on that side was longer than the duration of the displays on the other side. Similar findings were obtained when a sample of animals was tested for eye use during courtship displays in the presence of a female. Furthermore, consistency in eye use was observed in fish tested first with the mirror and then, 2 months later, with the female. Results are discussed with respect to the issue of the evolution of brain lateralization.

Lateralization of predator-evasion response in a teleost fish (Girardinus falcatus).

Evidence of lateral asymmetries in the direction of turning during escape behaviour in a species of poeciliid fish, Girardinus falcatus, is reported. When repeatedly faced with a simulated predator (in five successive sessions, spaced 7 days apart), immature Girardinus falcatus exhibited a significant population bias to turn right on the first session and a progressive bias to turn left in subsequent sessions. Mature Girardinus were then tested to check whether the shift in the direction of turn with repeated sessions depended on maturation or habituation. It was found that adult Girardinus showed a slight population bias to turn right in the first session and a strong subsequent bias to turn left after repeated sessions. The implications of these findings to our current understanding of the evolution of brain lateralization are discussed.