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Once the emperor Hui Tsung was enjoying the sight of a lichee tree laden with fruit before the palace when a peacock approached the tree, and he summoned his artists at once to make a picture. They produced a magnificent painting of the peacock with its right foot poised to take a step on a flower-bed: but to their surprise the emperor shook his head over it. A few days later when he asked if they had discovered their mistake, they had no answer ready. Then Hui Tsung told them: ‘A peacock always raises its left foot first to climb.’
Cheng Chen-To, Chang Heng and Hsu Pang-Ta (1957)
Summary
Once thought to be unique to the human brain, lateralization of structure and behaviour is now known to be widespread in vertebrates and, furthermore, it has a similar plan of organization in the different species. This chapter introduces the basic pattern of lateralization of vertebrate species and does so in a historical context to highlight the fact that, until some 20 years ago, it was widely and incorrectly assumed that having a lateralized brain was a mark of the cognitive superiority of humans. It also introduces some of the new evidence showing the presence of lateralization in invertebrate species.
Experience can enhance, suppress or change in other ways the development of lateralization. Exactly which of these occurs depends on the species, the nature of the experience and the stage of life at which it takes place. Lateralization of individuals and groups can be modulated by experience and by steroid hormones. The latter may be important in the development of sex differences in lateralization. Research in this area is in its infancy compared with our knowledge of species differences in lateralization, but we are able to give some potent examples to illustrate the importance of experience and hormone levels at particular stages of development.
Introduction
The brain is not as hard-wired as once thought. It changes its connections in response to experience, especially in early life but also in adulthood. Some regions of the brain even change size in response to specific kinds of experience. The hippocampus is such a region. In humans, we know that the hippocampus has a special role in spatial memory. A study of London taxi drivers has shown that they have a larger than average posterior region of the right hippocampus and a smaller than average anterior region of the hippocampus (Maguire et al., 2000). In animals too, the size of the hippocampus is related to spatial ability. Species that cache food and retrieve it at a later time have a larger hippocampus than do closely related non-caching species. This is known to be the case in squirrels (Johnson et al., 2010), kangaroo rats (Jacobs and Spencer, 1994) and several species of birds, including marsh tits and Clarke’s nutcrackers (Shettleworth, 2003).
Lateralization is manifested in two main ways: (1) in individuals but with no common direction (bias) in the group or population, or (2) in individuals and in the same direction in most individuals so that the group or population is biased. The first is discussed in terms of evidence of efficiency of neural processing in a lateralized brain. The second is discussed mainly in terms of the hypothesis that population biases occur as evolutionarily stable strategies when lateralized individuals coordinate with each other. This hypothesis is supported both by recent evidence and by mathematical models.
Introduction
It is believed that bilateral symmetry evolved when organisms adopted an axial orientation to their direction of movement and it is usually agreed that the pathway to a bilateral nervous system led from radial symmetry. In addition to being bilaterally divided, however, the nervous system of vertebrates shows a pervasively contralateral organization in that afferent and efferent pathways cross the midline of the body so that each side of the brain connects to the opposite side of the body. Also, as we have already seen (Chapter 1), the nervous system has a certain degree of asymmetry between the left and the right sides, and this is seen in both function and structure. Before considering the function of such an asymmetrical organization let us discuss the problem of why the nervous system is organized contralaterally (Figure 2.1), given that we shall refer to such an organization almost continuously while describing experiments and observations on asymmetries in animal behaviour.
This chapter discusses the evolution of lateralization in vertebrates and their ancestors. Vertebrate asymmetry was dominated from the start by extraordinary bodily asymmetry, which determined the course of evolution of nervous system asymmetries. Modern Echinoderms (starfish and sea urchins) and all chordates (the group to which vertebrates belong) came from an ancestor that had extreme right–left asymmetry. Evidence from invertebrates, discussed later in this chapter, suggests that the possession of paired sense organs (including sensory inputs from paired appendages, as well as from paired eyes) has sometimes been sufficient to allow the evolution of lateralization of functions of the central nervous system. The evidence takes us far from the earlier notion that hemispheric specialization evolved in humans about 2.5 million years ago along with language, handedness and tool using. Although important steps in human evolution, discussed here, involved brain lateralization, they were shaped by pre-existing asymmetries, rather than appearing de novo.
Origins of asymmetry in chordates
Ancestral chordates lived in a marine environment (in the Cambrian/Precambrian periods) very different from any present today. Food was available as tiny algae near the surface of the sea and as organic remains that had sunk to the sea floor; both were exploited by animals. The rarity of deep burrowing forms of life (Bambach et al., 2007) meant that accumulation of edible particles in the deposits on the sea floor was greater than now. The structure of food webs (‘what ate what’) reveals that, because efficient predators were relatively scarce (albeit not non-existent, as shown by the recent discovery of a Cambrian, arthropod predator; Paterson et al., 2011), many organisms did well at this time without the need for very effective defences against predators or the ability to flee (Bambach et al., 2007).
An inventory has been made of the decapod fauna of the Glorieuses Islands, western Indian Ocean (WIO), following the BIORECIE 2 Expedition to the Islands, 5–17 December 2012. Field data are complemented by a review of taxonomic studies for these islands. Overall 157 species are now reported from the Glorieuses Islands, including 61 new records. The presence of the coconut crab, Birgus latro, is confirmed from these islands, for the first time since 1884, and the WIO distribution of this endangered species is updated, based on observations made in the region since 2006.
Three species of the serpulid genus Hydroides are described and illustrated herein. Hydroides bannerorum and H. albiceps are newly recorded to Guam in the Mariana Islands. Hydroides perezi is also described and illustrated from specimens collected in Pago Bay. We provide notes on the opercular ontogenetic variation of H. albiceps to help clarify possible confusions with opercula of similar taxa.
This paper reports the first record of the occurrence of Pinna fragilis from Korangi Creek, Pakistan coast (24°47′N 67°11′E). A description of the species collected from the Pakistan coast is given along with size, habitat, ecology and economic importance. Shells of 114–347 mm were found from the area. The dark grey pearls of dull lustre were found from few shells.
Cylichna villersii, a minute ‘bubble snail’ is newly recorded in the Mediterranean Sea from specimens recently collected along the Israeli coastline. The species, described and illustrated, matches the syntype collected in the Red Sea by Savigny. It is the eighth Erythraean alien cephalaspideid species recorded in the Levantine Basin. The recent collection of many living specimens in several continuously sampled locations attests to the speed of its establishment in the south-eastern Levant.
We present a species checklist, characterizing the ichthyofaunal composition of four coastal archipelagos in waters off the metropolitan area of Rio de Janeiro, Brazil. Data were obtained from ~120 h of underwater observations, published and unpublished information, including new species records. A total of 197 fish species (78 new records), belonging to 130 genera and 61 families, were observed. The family Carangidae (17) dominated in number of species, followed by Labridae and Serranidae (10 each), Epinephelidae and Scaridae (9 each). Most species are distributed along the tropical western Atlantic (51.8%) or on both sides of the Atlantic Ocean (20.5%). A minor part is endemic to Brazil (9.7%) or range to temperate rocky reefs of Patagonia (6.7%). Fourteen species are threatened, according to the IUCN, and listed under the Brazilian Ministry of the Environment red list (6.6%). Despite an increment of fish species listed herein, and the recent establishment/creation of a Marine Protected Area (Monumento Natural das Ilhas Cagarras—MoNa Cagarras), research efforts are clearly needed along these archipelagos, as nearshore environmental threats increase due to lack of regulatory practices.
Carcasses of whales provide much valuable information on their natural history. However, some specimens cannot be identified in the field due to the advanced state of decomposition. In this study, the DNA was extracted and the mitochondrial cytochrome b gene was amplified by polymerase chain reaction and sequenced for four carcasses of possible mysticeti (GEMM: 075, 088, 135 and GEMARS: 1302). A blast search using the nucleotide–nucleotide basic local alignment (blastn) search tool was conducted using the generated sequences. Samples GEMM 075 and GEMARS 1302 showed 98% identity to one sequence of Balaenoptera acutorostrata. Samples GEMM 088 and GEMM 135 showed 99% identity to sequences from Balaenoptera edeni and Megaptera novaeangliae, respectively. A neighbour-joining tree was generated using sequences from GenBank from all species of balaenopterid that occur on the coast of Brazil. The results showed that all carcasses analysed were correspondent to species from the family Balaenopteridae already recorded in Brazil.
This work provides the first information about the stomach contents of juvenile green turtles (Chelonia mydas) on the coast of Rio de Janeiro State, south-eastern Brazil (22°50′S–23°00′S). Between June 2009 and May 2010, we obtained 49 stomach contents of specimens that were stranded, of which 37 contained food items. Among food items, macroalgae stood out with 83.7% frequency of occurrence in the stomachs analysed and 37 species were identified. The most representative macroalgae in the stomach contents were: Sargassum vulgare, Ulva lactuca, Gelidiella acerosa and Pterocladiella capillacea. Teleosts and molluscs were also recorded in the stomach contents of the species in the region; however, with low representativeness. Analysis of stomach contents confirmed herbivory as the preferred dietary pattern of C. mydas.
The Atlantic spotted dolphin, Stenella frontalis, presents two geographically isolated populations in the western South Atlantic. This note reports on a 350 km northward extension range of the southern population. The sighted group of 80 animals was in waters 66 m deep and 75 nautical miles distant from the coast. The record was observed during a dedicated cetacean survey in Brazilian waters in the spring of 2008 and supports a discontinuous distribution along the coast of Brazil.
The presence of the pantropical ascidian Microcosmus exasperatus has been an object of some taxonomic confusion in the Mediterranean Sea. A closely related species, M. squamiger was previously reported in the north of Tunisia (in 1962) as M. exasperatus; it has spread to the western Mediterranean and the northern part of the Ionian Sea, whereas, the reliable report of M. exasperatus has been made more recently (in 1998) in the Gulf of Gabès (South Tunisia), and later studies have confirmed its presence in the eastern Mediterranean basin (Lebanon and Israel). The occurrence of this species in Izmir (Turkey) extends its distribution range to the Aegean Sea. At present, the distributions of both species of Microcosmus are not overlapping in the Mediterranean, suggesting a different route of entrance, probably by the maritime traffic (M. squamiger via the Strait of Gibraltar; and M. exasperatus via the Suez Canal). The discontinuity in the distribution of M. exasperatus in the Mediterranean is analysed and discussed.
Four porcellanid species, Petrolisthes coccineus (Owen, 1839), P. lamarckii (Leach, 1820), P. moluccensis (De Man, 1888) and P. tomentosus (Dana, 1852), are reported on the basis of specimens collected from an intertidal area of Agatti Island, Lakshadweep Archipelago in south-western India. All the four species are recorded from the archipelago for the first time, and P. moluccensis is new to the Indian fauna. Morphological diagnoses, notes on the habitats and distributions are provided for the four species.
Two shark species, Odontaspis ferox and Centrophorus squamosus, are recorded for the first time at the Galápagos Islands based on one sighting reported by tourist divers and one specimen captured by a local fisherman. The identification of a third shark species, Mustelus albipinnis, previously unidentified and recorded as Mustelus sp., is achieved based on several sightings at the same location.
The type species of Licmophora, L. flabellata (Carm.) C. Ag., is atypical for the genus in having a row of intercalary rimoportulae along the sternum and four plate-like plastids with pyrenoids. The pattern of rimoportulae is sufficiently unique to have prompted the question of whether this species should be separated from the genus. We report here L. comnavmaria sp. nov., which also has a row of intercalary rimoportulae. Its plastids are numerous small plates with a pyrenoid, different from plastids in both L. flabellata and other Licmophora species. Like L. flabellata, the new species forms extensive colonies on long, branched, multistranded mucilage stalks. The new species is distinguished from L. flabellata also by the delicate silicification and the extremely fine striae (55 vs 30–35 in 10 µm). Only one other species, L. pfannkucheae Giffen, has been described with intercalary rimoportulae (plastids unknown) and that species may not be distinct from L. flabellata. Thus the present species is a significant addition to the genus.
Delicate shallow-water sea pens were collected from Mangamaripeta fish landing centre of Visakhapatnam, India where they were brought in along with the fish catch. The sea pens were preserved and identified as Cavernularia obesa Valenciennes in Milne Edward & Haime, 1850. The species is being recorded for the first time from these waters.