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Clinical Biochemistry of Domestic Animals (Sixth Edition) - UMK ...

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562<br />

Chapter | 18 Pituitary Function<br />

The neurosecretory pathway runs from the anterior<br />

hypothalamus, traverses the floor <strong>of</strong> the ventral hypothalamus,<br />

and terminates in the neural lobe (NL) <strong>of</strong> the neurohypophysis<br />

on fenestrated blood vessels ( Page, 1986 ). The<br />

system is involved in osmoregulation through the production<br />

and release <strong>of</strong> vasopressin, and in parturition and nursing<br />

through the secretion <strong>of</strong> oxytocin.<br />

In the intermediate lobe (IL) the secretory activity is regulated<br />

via direct neuronal inhibitory and stimulatory influences<br />

( Saland, 2001 ). In amphibians ( Vazquez-Martinez et al. ,<br />

2003 ) and reptiles ( Dores et al. , 1987 ), the IL plays an<br />

important role in adaptation to background color. The function<br />

<strong>of</strong> IL cells in mammals has not been fully established,<br />

but they may play a role in opioid-regulated functions.<br />

A . Anatomical Considerations<br />

The hypothalamus and pituitary control vital functions<br />

such as growth, reproduction, lactation, basal metabolism,<br />

stress response, parameters <strong>of</strong> immune function, and the<br />

state <strong>of</strong> hydration. Understanding <strong>of</strong> the complicated functional<br />

relationship <strong>of</strong> the hypothalamus to the pituitary<br />

requires an appreciation <strong>of</strong> the anatomical relationships.<br />

1 . Hypothalamus<br />

Hypophysiotropic neurohormones are produced in several<br />

areas <strong>of</strong> the hypothalamus. For example, in an immun<strong>of</strong>luorescence<br />

study <strong>of</strong> hypothalami <strong>of</strong> dogs, the majority <strong>of</strong><br />

cell bodies with immunoreactivity to corticotropin-releasing<br />

hormone (CRH) was found in the region <strong>of</strong> the periventricular<br />

and paraventricular nuclei, but they were also<br />

found in the supraoptic and suprachiasmatic area as well as<br />

craniodorsal to the mammillary bodies ( Stolp et al. , 1987 ).<br />

The cell bodies <strong>of</strong> the neurohormone-producing neurons<br />

that project to the median eminence are in part<br />

intermingled with cell bodies that also synthesize these<br />

neurohormones but project to other brain areas. The majority<br />

<strong>of</strong> the neurons that project to the median eminence are<br />

found in the preoptic and suprachiasmatic region <strong>of</strong> the<br />

hypothalamus. Axons containing the same neurohormone<br />

may have synaptic contacts that enable regulation <strong>of</strong> cellular<br />

function between these neurons.<br />

The neurohormone-producing cells receive a complex<br />

neural input from a variety <strong>of</strong> chemical messengers, such as<br />

neurotransmitters and other neurohormones. Not only the<br />

neurohormones CRH and arginine vasopressin (AVP), but<br />

in general the combination <strong>of</strong> a neurohormone and another<br />

chemical messenger, may colocalize within a single neuron.<br />

The neurons <strong>of</strong> the neurohypophyseal system represent<br />

a more anatomically distinct entity, with cell bodies<br />

located in the paraventricular and supraoptic hypothalamic<br />

nuclei ( Sawchenko and Swanson, 1983 ). However, within<br />

these areas there are also neurons producing a variety <strong>of</strong><br />

other neuropeptides.<br />

2 . Neurotransmitter Systems<br />

The major neurotransmitter systems for intercellular communication<br />

within the central nervous system consist <strong>of</strong><br />

monoamines and peptides. These chemical messengers<br />

regulate the biosynthesis and release <strong>of</strong> the hypophysiotropic<br />

neurohormones. Through a network <strong>of</strong> axodendritic<br />

and axoaxonic contacts, these neurons are connected to the<br />

neurohormone-producing cells. In addition, many monoamines<br />

and peptides are found within the hypophysiotropic<br />

hormone-producing cells, where they are released together<br />

with the neurohormones into the portal system and modify<br />

the effect <strong>of</strong> hypothalamic hormones on the pituitary.<br />

The biogenic amine neurotransmitters, known to play<br />

a regulating/modulating role in the hypothalamic-pituitary<br />

system, include catecholamines (dopamine, noradrenalin,<br />

and adrenalin), indolamines (serotonin, melatonin), acetylcholine,<br />

γ -aminobutyric acid (GABA), and histamine.<br />

Neuropharmacological agents can be used to alter neurotransmitter<br />

effects and, as a consequence, hypothalamic<br />

and pituitary hormone release.<br />

Many <strong>of</strong> the peptides with potential effects on hypophysiotropic<br />

hormone release are widely distributed in<br />

hypothalamic and extrahypothalamic areas <strong>of</strong> the brain.<br />

They include, among many others, peptides common to the<br />

gastrointestinal tract, such as gastrin, cholecystokinin, and<br />

pancreatic polypeptide as well as bombesin, angiotensin II,<br />

galanin, substance P, neurotensin, enkephalins, neuropeptide<br />

Y, natriuretic peptide, the vasoactive intestinal peptide<br />

(VIP), and the peptide histidine isoleucine (PHI). The last<br />

three peptides may, through vasoconstriction and vasodilatation<br />

activities, play an important role in the control <strong>of</strong> the<br />

portal blood flow.<br />

3 . Vascular System<br />

The releasing and inhibiting hormones are stored in nerve<br />

terminals in the median eminence, where their concentrations<br />

are 10 to 100 times as great as elsewhere in the hypothalamus.<br />

The uniquely organized capillary plexus ( Halasz,<br />

1994 ; Page, 1986 ) <strong>of</strong> the median eminence is in close proximity<br />

to nerve terminals <strong>of</strong> the hypophysiotropic neurons.<br />

In contrast to other brain regions, the blood-brain barrier<br />

in the area <strong>of</strong> the median eminence is incomplete, permitting<br />

protein and peptide hormones as well as other charged<br />

particles to move to the intercapillary spaces and the nerve<br />

terminals contained therein. These terminals respond to<br />

humoral and neuronal stimuli by secreting releasing and<br />

inhibiting factors into the portal system.<br />

The portal capillaries coalesce into a series <strong>of</strong> vessels<br />

that descend through the pituitary stalk and form a second<br />

capillary plexus that surrounds the AL cells ( Fig. 18-1 ).<br />

Inferior hypophyseal arteries supply the neurohypophysis.<br />

From the primary plexus <strong>of</strong> the neural lobe (NL), blood<br />

flows not only to the systemic circulation but also to the AL<br />

and the hypothalamus. There is evidence for some degree <strong>of</strong>

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