The most striking, interesting, yet poorly understood gross morphological features of the cerebral hemispheres in mammals are the diverse and complex arrangements of their cortical gyri and sulci (Fig. 1). Among mammals, the spinal cord and brain-stem nuclei are morphologically quite similar, despite variations in size. However, during evolution, the cerebrum and cerebellum have undergone marked variations in size, shape, and convolutional complexity (Fig. 2). External morphological features of mammalian brains have long been utilized to judge not only the degree of phylogenetic development, but also the nature and level of complexity of brain functions. The great variety of living mammals exhibit a corresponding variety of brain shapes, sizes, and patterns of fissuration and convolution of the cerebral neocortex. The view has consistently been expressed that animals with brains having greater amounts of convoluted cerebral neocortex were more intelligent as well as perceptually and behaviorally more complex.

Why Does Cerebral Cortex Fissure and Fold

The present investigation was designed to determine the number and internal organization of somatosensory fields in monotremes. Microelectrode mapping methods were used in conjunction with cytochrome oxidase and myelin staining to reveal subdivisions and topography of somatosensory cortex in the platypus and the short-billed echidna. The neocortices of both monotremes were found to contain four representations of the body surface. A large area that contained neurons predominantly responsive to cutaneous stimulation of the contralateral body surface was identified as the primary somatosensory area (SI). Although the overall organization of SI was similar in both mammals, the platypus had a relatively larger representation of the bill. Furthermore, some of the neurons in the bill representation of SI were also responsive to low amplitude electrical stimulation. These neurons were spatially segregated from neurons responsive to pure mechanosensory stimulation. Another somatosensory field (R) was identified immediately rostral to SI. The topographic organization of R was similar to that found in SI; however, neurons in R responded most often to light pressure and taps to peripheral body parts. Neurons in cortex rostral to R were responsive to manipulation of joints and hard taps to the body. We termed this field the manipulation field (M). The mediolateral sequence of representation in M was similar to that of both SI and R, but was topographically less precise. Another somatosensory field, caudal to SI, was adjacent to SI laterally at the representation of the face, but medially was separated from SI by auditory cortex. Its position relative to SI and auditory cortex, and its topographic organization led us to hypothesize that this caudal field may be homologous to the parietal ventral area (PV) as described in other mammals. The evidence for the existence of four separate representations in somatosensory cortex in the two species of monotremes indicates that cortical organization is more complex in these mammals than was previously thought. Because the two monotreme families have been separate for at least 55 million years (Richardson, B.J. [1987] Aust. Mammal. 11:71-73), the present results suggest either that the original differentiation of fields occurred very early in mammalian evolution or that the potential for differentiation of somatosensory cortex into multiple fields is highly constrained in evolution, so that both species arrived at the same solution independently.

Organization of somatosensory cortex in monotremes: in search of the prototypical plan.

Latex endocranial casts, which may be made without damaging the skull, reveal much of the information provided by a hemisected skull. Examination of drawings of endocasts superimposed on skulls may provide insight into the biological significance of skull and brain morphology. The high degree of cranial flexion and the globose brain shape of Daubentonia appear to be related to the functional demands of its gnawing mechanism. The broad frontal lobes of indriids are correlated with orbital orientation; differences in frontal lobe sulcal pattern suggest greater elaboration of the motor filed for the hand in indriids than in lemurids. Several features of lorisid cranial anatomy are discussed. It is suggested that, as a first approximation, increased splanchnocranial declination in small prosimians results from the necessity of accommodating relatively large eyes in a skull with a relatively small splanchnocranium.

A new approach to mammalian cranial analysis, illustrated by examples of prosimian primates

We review the evidence for the concept of the “initial” or prototype brain. We outline four possible modes of brain evolution suggested by our new findings on the evolutionary status of the dolphin brain. The four modes involve various forms of deviation from and conformity to the hypothesized initial brain type. These include examples of conservative evolution, progressive evolution, and combinations of the two in which features of one or the other become dominant. The four types of neocortical organization in extant mammals may be the result of selective pressures on sensory/motor systems resulting in divergent patterns of brain phylogenesis. A modular “modification/multiplication” hypothesis is proposed as a mechanism of neocortical evolution in eutherians. Representative models of the initial ancestral group of mammals include not only extant basal Insectivora but also Chiroptera; we have found that dolphins and large whales have also retained many features of the archetypal or initial brain. This group evolved from the initial mammalian stock and returned to the aquatic environment some 50 million years ago. This unique experiment of nature shows the effects of radical changes in environment on brain-body adaptations and specializations. Although the dolphin brain has certain quantitative characteristics of the evolutionary changes seen in the higher terrestrial mammals, it has also retained many of the conservative structural features of the initial brain. Its neocortical organization is accordingly different, largely in a quantitative sense, from that of terrestrial models of the initial brain such as the hedgehog.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0140525X0005281X

Implications of the “initial brain” concept for brain evolution in Cetacea

Sensory and motor representation in the cerebral cortex of the three-toed sloth (Bradypus tridactylus).

Sensory projections of somatic, auditory and visual origin to the cerebral cortex of the sloth Choloepus hoffmanni Peters).

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Sensory projections of somatic, auditory and visual origin to the cerebral cortex of the sloth Choloepus hoffmanni Peters).