adrenal gland

Development

The adrenal gland forms from two primordia: cells of mesodermal origin which give rise to the steroid-secreting cells, and neural cells of ectodermal origin which develop into the catecholamine-secreting tissue (also known as chromaffin tissue). In higher vertebrates, mesenchymal cells originating from the coelomic cavity near the genital ridge proliferate to form a cluster of cells destined to be the adrenal cortex. During the second month of human development, cells of the neural crest migrate to the region of the developing adrenal and begin to proliferate on its surface. The expanding cortical tissue encapsulates the neural cells forming the cortex and medulla. In mammals, three distinct zones form within the cortex: the outermost zona glomerulosa, the middle zona fasiculata, and the inner zona reticularis. The glomerulosa cells contain an enzyme, aldosterone synthase, which converts corticosterone to aldosterone, the principal steroid (mineralocorticoid) secreted from this zone. The inner zones (fasiculata and reticularis) primarily secrete glucocorticoids and large amounts of sex steroid precursors. In many lower vertebrates, the two tissues form from similar primordia but migrate and associate in different ways to the extent that in some cases the two tissues develop in isolation from each other.

Comparative anatomy

While the paired adrenals in mammals have a characteristic cortical-medullary arrangement with distinct zonation present in the cortex, such distinctions are lacking in nonmammalian species. In more primitive fishes, chromaffin cells form in isolation from steroidogenic tissue. A general trend is present, however, throughout vertebrates for a closer association of chromaffin and steroidogenic tissues. Zonation in steroidogenic tissue is largely confined to mammals, although suggestions of separate cell types have been postulated in birds and in some other species.

Comparative endocrinology

Hormones are secreted from the cells of both the medulla and the cortex.

Chromaffin cells

In all vertebrates, chromaffin cells secrete catecholamines into circulation. In most species, the major catecholamine secreted is epinephrine, although significant amounts of norepinephrine are released by many animals. Some dopamine is also secreted. No phylogenetic trend is obvious to explain or predict the ratio of epinephrine to norepinephrine secreted in a given species. A given species may release the two catecholamines in different ratios, depending on the nature of the stimulus. The great majority of the norepinephrine in circulation actually originates from that which is released from non-adrenal sympathetic nerve endings and leaks into the bloodstream. In addition to catecholamines, chromaffin cells secrete an array of other substances, including proteins such as chromogranin A and opioid peptides. See Epinephrine

Biologic effects of catecholamines are mediated through their binding to two receptor classes, α- and β-adrenergic receptors. Further examination of these receptors has revealed that subclasses of each type exist and likely account for the responses on different target tissues. In general, biologic responses to catecholamines include mobilization of glucose from liver and muscle, increased alertness, increased heart rate, and stimulation of metabolic rate.

Steroid hormones

In broad terms, most steroids secreted by adrenal steroidogenic cells are glucocorticoids, mineralocorticoids, or sex hormone precursors. However, these classes have been established largely on the basis of differential actions in mammals. The principal glucocorticoids are cortisol and corticosterone, while the main mineralocorticoid is aldosterone. This division of action holds for mammalian species and likely for reptiles and birds. In other vertebrates, such as fish and amphibians, steroids from the interrenal tissue do not show such specialized actions; instead, most show activities of both glucocorticoid and mineralocorticoid type. Mammals, birds, reptiles, and amphibians secrete cortisol, corticosterone, and aldosterone. The ratios of the two glucocorticoids vary across species; in general, corticosterone is the more important product in nonmammalian species. Even within mammals, a large variation exists across species, due to the relative ratio of cortisol to corticosterone from the adrenal cortex.

Effects of adrenal-derived steroids in lower vertebrates involve a diverse array of actions, including control of distribution and availability of metabolic fuels such as glucose, and regulation of sodium and extracellular fluid volume. In nonmammalian vertebrates, corticosterone, cortisol, and aldosterone possess mineralocorticoid effects. Other areas where adrenal steroids likely contribute to biologic processes include control of protein, fat, and carbohydrate balance; reproduction; and growth and development. See Steroid