The olfactory system of mammals can be divided into a main and accessory olfactory system with initial processing for each system occurring in the olfactory bulb. The main and accessory olfactory bulbs have similar structural features, even though they appear to be functionally independent. In mammals the main olfactory bulb (MOB) is also one of two established sites of lifelong generation of new cells. The present study describes the histological and immunohistochemical neuroanatomy of the olfactory bulb of the African elephant (Loxodonta africana). The morphology of MOB of the elephant does not differ significantly from that described in other mammals; however, it lacks the internal plexiform layer. In addition, the glomeruli of the glomerular layer are organised in 2-4 "honey-combed" layers, a feature not commonly observed. The cell types and structures revealed with immunohistochemical stains (parvalbumin, calbindin, calretinin, tyrosine hydroxylase, orexin-A, glial fibrillary acidic protein) were similar to other mammals. Neurogenesis was examined using the neurogenic marker doublecortin. Migration of newly generated cells was observed in most layers of the MOB. No accessory olfactory bulb (AOB) was observed. Based on the general anatomy and the immunohistochemical observations, it is evident that the morphology of the African elephant MOB is, for the most part, similar to that of all mammals, although very large in absolute size.

It is a central assumption that larger bodies require larger brains, across species but also possibly within species with continuous growth throughout the lifetime, such as the crocodile. The current study investigates the relationships between body growth (length and mass) and the rates of growth of various subdivisions of the central nervous system (CNS) (brain, spinal cord, eyes) in Nile crocodiles weighing between 90 g and 90 kg. Although the brain appears to grow in two phases in relation to body mass, initially very rapidly then very slowly, it turns out that brain mass increases continuously as a power function of body mass with a small exponent of 0.256, such that a 10-fold increase in body mass is accompanied by a 1.8-fold in brain mass. Eye volume increases slowly with increasing body mass, as a power function of the latter with an exponent of 0.37. The spinal cord, however, grows more rapidly in mass, accompanying body mass raised to an exponent of 0.54, and increasing in length as predicted, with body mass raised to an exponent of 0.32 (close to the predicted 1/3). While supporting the expectation formulated by Jerison that larger bodies require larger brains to operate them, our findings show that: (1) the rate of increase in brain size is very small compared to body growth; and (2) different parts of the CNS grow at different rates accompanying continuous body growth, with a faster increase in spinal cord mass and eye volume, than in brain mass. Anat Rec, 296:1489-1500, 2013. V C 2013 Wiley Periodicals, Inc.

It is generally believed that animals with larger bodies require larger brains, composed of more neurons. Across mammalian species, there is a correlation between body mass and the number of brain neurons, albeit with low allometric exponents. If larger bodies imperatively require more neurons to operate them, then such an increase in the number of neurons should be detected across individuals of a continuously growing species, such as the Nile crocodile. In the current study we use the isotropic fractionator method of cell counting to determine how the number of neurons and non-neurons in 6 specific brain regions and the spinal cord change with increasing body mass in the Nile crocodile. The central nervous system (CNS) structures examined all increase in mass as a function of body mass, with allometric exponents of around 0.2, except for the spinal cord, which increases with an exponent of 0.6. We find that numbers of non-neurons increase slowly, but significantly, in all CNS structures, scaling as a function of body mass with exponents ranging between 0.1 and 0.3. In contrast, numbers of neurons scale with body mass in the spinal cord, olfactory bulb, cerebellum and telencephalon, with exponents of between 0.08 and 0.20, but not in the brainstem and diencephalon, the brain structures that receive inputs and send outputs to the growing body. Densities of both neurons and non-neurons decrease with increasing body mass. These results indicate that increasing body mass with growth in the Nile crocodile is associated with a general addition of non-neurons and increasing cell size throughout CNS structures, but is only associated with an addition of neurons in some structures (and at very small rates) and not in those brain structures directly connected to the body. Larger bodies thus do not imperatively require more neurons to operate them.

The brain of the crocodile is known to gain in mass allometrically throughout life, and the addition of neurons (as well as non-neurons) appears to play a significant role in this increasing brain mass. We used immunohistochemistry in the brains of 12 Nile crocodiles ranging between 350 g and 86 kg in body mass and 1.99 g to 7.9 g in brain mass to identify the regions of the brain in which neurons immunopositive for doublecortin (DCX), a marker for potential adult neurogenesis, are found. Similar to other reptiles, potential newly born neurons, those immunopositive for DCX, were found throughout the telencephalon, the main and accessory olfactory bulbs and the olfactory tract, and in the cerebellar cortex; however, no DCX immunopositive neurons were observed in the diencephalon or brainstem. An apparent moderate decrease in the density of DCX labeled neurons in the olfactory bulbs and tract as well as the cerebellar cortex was observed with increasing brain mass, but the observed qualitative density of labeled neurons within the telencephalon was maintained irrespective of brain mass. Three potential neurogenic zones, within the sulci of the lateral ventricle, were identified, and these are similar to those seen in other reptiles. This study indicates that at least part of the gain in brain mass with age in the Nile crocodile may be accounted for by the potential addition and integration of new neurons into the existing circuitry, especially so for the olfactory system, telencephalon and cerebellar cortex. Anat Rec, 301:659-672, 2018. © 2017 Wiley Periodicals, Inc.

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