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Chapter 18 Pituitary Function Jan A. Mol Department of Clinical Sciences of Companion Animals Faculty of Veterinary Medicine Utrecht University Utrecht, The Netherlands Björn P. Meij Department of Clinical Sciences of Companion Animals Faculty of Veterinary Medicine Utrecht University Utrecht, The Netherlands I. HYPOTHALAMUS-PITUITARY SYSTEM A. Anatomical Considerations B. Regulation of Pituitary Functions II. ANTERIOR LOBE AND INTERMEDIATE LOBE A. Proopiomelanocortin-Derived Peptides B. Glycoprotein Hormones C. Somatomammotropic Hormones III. NEUROHYPOPHYSIS A. Vasopressin B. Oxytocin IV. ASSESSMENT OF PITUITARY FUNCTION A. Adenohypophysis B. Neurohypophysis REFERENCES I . HYPOTHALAMUS-PITUITARY SYSTEM The hypothalamic-pituitary system is a preeminent example of the integration of neural and endocrine control. It consists of three major systems: (1) a neuroendocrine system connected to an endocrine system by a portal circulation, (2) a neurosecretory pathway, and (3) a direct neural regulation of endocrine secretion ( Fig. 18-1 ). The neuroendocrine system involves clusters of peptideand monoamine-secreting cells in the anterior and midportion of the ventral hypothalamus. Their products reach the median eminence by axonal transport. From there they are released into the capillary vessels of the hypothalamicpituitary portal system and transported to the pituitary to regulate the secretion of hormones from the anterior lobe (AL) of the adenohypophysis ( Amar and Weiss, 2003 ). Hypophysiotropic hormones GHRH TRH GnRH CRH Others IL NL FIGURE 18-1 Schematic representation of the relationship between the hypothalamus and the pituitary in a generalized mammal. The hypothalamus exerts control over the anterior lobe (AL) by releasing and inhibiting factors. These hormones reach the AL cells via capillaries of the pituitary portal system. The neural lobe (NL) of the pituitary is a downward projection of the hypothalamus. The intermediate lobe (IL) is under direct neurotransmitter control. AL Pituitary hormones ACTH GH PRL TSH LH FSH α MSH VP OT Clinical Biochemistry of Domestic Animals, 6th Edition 561 Copyright © 2008, Elsevier Inc. All rights reserved.
562 Chapter | 18 Pituitary Function The neurosecretory pathway runs from the anterior hypothalamus, traverses the floor of the ventral hypothalamus, and terminates in the neural lobe (NL) of the neurohypophysis on fenestrated blood vessels ( Page, 1986 ). The system is involved in osmoregulation through the production and release of vasopressin, and in parturition and nursing through the secretion of oxytocin. In the intermediate lobe (IL) the secretory activity is regulated via direct neuronal inhibitory and stimulatory influences ( Saland, 2001 ). In amphibians ( Vazquez-Martinez et al. , 2003 ) and reptiles ( Dores et al. , 1987 ), the IL plays an important role in adaptation to background color. The function of IL cells in mammals has not been fully established, but they may play a role in opioid-regulated functions. A . Anatomical Considerations The hypothalamus and pituitary control vital functions such as growth, reproduction, lactation, basal metabolism, stress response, parameters of immune function, and the state of hydration. Understanding of the complicated functional relationship of the hypothalamus to the pituitary requires an appreciation of the anatomical relationships. 1 . Hypothalamus Hypophysiotropic neurohormones are produced in several areas of the hypothalamus. For example, in an immunofluorescence study of hypothalami of dogs, the majority of cell bodies with immunoreactivity to corticotropin-releasing hormone (CRH) was found in the region of the periventricular and paraventricular nuclei, but they were also found in the supraoptic and suprachiasmatic area as well as craniodorsal to the mammillary bodies ( Stolp et al. , 1987 ). The cell bodies of the neurohormone-producing neurons that project to the median eminence are in part intermingled with cell bodies that also synthesize these neurohormones but project to other brain areas. The majority of the neurons that project to the median eminence are found in the preoptic and suprachiasmatic region of the hypothalamus. Axons containing the same neurohormone may have synaptic contacts that enable regulation of cellular function between these neurons. The neurohormone-producing cells receive a complex neural input from a variety of chemical messengers, such as neurotransmitters and other neurohormones. Not only the neurohormones CRH and arginine vasopressin (AVP), but in general the combination of a neurohormone and another chemical messenger, may colocalize within a single neuron. The neurons of the neurohypophyseal system represent a more anatomically distinct entity, with cell bodies located in the paraventricular and supraoptic hypothalamic nuclei ( Sawchenko and Swanson, 1983 ). However, within these areas there are also neurons producing a variety of other neuropeptides. 2 . Neurotransmitter Systems The major neurotransmitter systems for intercellular communication within the central nervous system consist of monoamines and peptides. These chemical messengers regulate the biosynthesis and release of the hypophysiotropic neurohormones. Through a network of axodendritic and axoaxonic contacts, these neurons are connected to the neurohormone-producing cells. In addition, many monoamines and peptides are found within the hypophysiotropic hormone-producing cells, where they are released together with the neurohormones into the portal system and modify the effect of hypothalamic hormones on the pituitary. The biogenic amine neurotransmitters, known to play a regulating/modulating role in the hypothalamic-pituitary system, include catecholamines (dopamine, noradrenalin, and adrenalin), indolamines (serotonin, melatonin), acetylcholine, γ -aminobutyric acid (GABA), and histamine. Neuropharmacological agents can be used to alter neurotransmitter effects and, as a consequence, hypothalamic and pituitary hormone release. Many of the peptides with potential effects on hypophysiotropic hormone release are widely distributed in hypothalamic and extrahypothalamic areas of the brain. They include, among many others, peptides common to the gastrointestinal tract, such as gastrin, cholecystokinin, and pancreatic polypeptide as well as bombesin, angiotensin II, galanin, substance P, neurotensin, enkephalins, neuropeptide Y, natriuretic peptide, the vasoactive intestinal peptide (VIP), and the peptide histidine isoleucine (PHI). The last three peptides may, through vasoconstriction and vasodilatation activities, play an important role in the control of the portal blood flow. 3 . Vascular System The releasing and inhibiting hormones are stored in nerve terminals in the median eminence, where their concentrations are 10 to 100 times as great as elsewhere in the hypothalamus. The uniquely organized capillary plexus ( Halasz, 1994 ; Page, 1986 ) of the median eminence is in close proximity to nerve terminals of the hypophysiotropic neurons. In contrast to other brain regions, the blood-brain barrier in the area of the median eminence is incomplete, permitting protein and peptide hormones as well as other charged particles to move to the intercapillary spaces and the nerve terminals contained therein. These terminals respond to humoral and neuronal stimuli by secreting releasing and inhibiting factors into the portal system. The portal capillaries coalesce into a series of vessels that descend through the pituitary stalk and form a second capillary plexus that surrounds the AL cells ( Fig. 18-1 ). Inferior hypophyseal arteries supply the neurohypophysis. From the primary plexus of the neural lobe (NL), blood flows not only to the systemic circulation but also to the AL and the hypothalamus. There is evidence for some degree of
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Preface It has been more than a dec
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viii Contributors Björn P. Meij (5
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2 Chapter | 1 Concepts of Normality
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10 Chapter | 1 Concepts of Normalit
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Chapter 2 Comparative Medical Genet
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I. Introduction 29 Green Red Green
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I. Introduction 31 A1 genetic map d
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I. Introduction 33 of genomes of va
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36 Chapter | 2 Comparative Medical
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46 Chapter | 3 Carbohydrate Metabol
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IX. Disorders of Carbohydrate Metab
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X. Disorders of Ruminants Associate
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X. Disorders of Ruminants Associate
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References 79 Kaneko , J. J. , Luic
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Chapter 4 Lipids and Ketones Michae
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II. Long Chain Fatty Acids 83 FIGUR
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II. Long Chain Fatty Acids 85 Plasm
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IV. Phospholipids 87 The regulation
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V. Cholesterol 89 V. CHOLESTEROL A.
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VI. Lipoproteins 91 TABLE 4-1 Compo
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VI. Lipoproteins 93 TABLE 4-2 Apoli
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96 Chapter | 4 Lipids and Ketones B
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98 Chapter | 4 Lipids and Ketones
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Chapter 5 Proteins, Proteomics, and
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III. Metabolism of Proteins 119 C .
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IV. Plasma Proteins 121 NH 4 HCO
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V. Methodology 123 molecule. The gl
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V. Methodology 125 has occurred wil
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V. Methodology 127 Although attempt
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V. Methodology 129 kD 94 67 43 2 2
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V. Methodology 131 D . Specific Pro
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VI. Normal Plasma and Serum Protein
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VII. Interpretation of Serum Protei
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References 149 Andersson , M. , Ste
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References 151 response of haptoglo
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References 153 dogs with metastasiz
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References 155 Watson , T. D. G. (
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158 Chapter | 6 Clinical Veterinary
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V. Methods for Evaluation of the Im
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VI. Laboratory Diagnosis of Disease
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Chapter 7 The Erythrocyte: Physiolo
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II. Hematopoiesis 175 progenitor ce
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III. Developing Erythroid Cells 177
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IV. Mature RBC 185 immune-mediated
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IV. Mature RBC 187 TABLE 7-3 Erythr
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IV. Mature RBC 189 TABLE 7.5 Erythr
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IV. Mature RBC 191 Echinocyte forma
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IV. Mature RBC 193 Pig RBC isoantig
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IV. Mature RBC 195 occurs in chicke
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IV. Mature RBC 197 inorganic phosph
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IV. Mature RBC 199 (a) FIGURE 7-6 T
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IV. Mature RBC 201 RBC 2,3DPG incre
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IV. Mature RBC 203 mechanisms would
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IV. Mature RBC 205 released during
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IV. Mature RBC 207 reduced by ascor
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V. Determinants of RBC Survival 209
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V. Determinants of RBC Survival 211
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VI. Inherited Disorders of RBCs 213
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described in people. They include a
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References 221 Boas , F. E. , Forma
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References 223 Della Rovere , F. ,
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References 239 Williams , D. M. , L
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Chapter 8 Porphyrins and the Porphy
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II. Porphyrins 243 two vinyl groups
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II. Porphyrins 245 TABLE 8-2 Nomenc
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IV. Porphyrias 247 6. Uroporphyrins
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IV. Porphyrias 249 i . Urine It is
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IV. Porphyrias 251 to localize the
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IV. Porphyrias 253 FIGURE 8-6 Metab
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IV. Porphyrias 255 estrogens. The d
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References 257 and hexachlorobenzen
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Chapter 9 Iron Metabolism and Its D
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II. Iron Distribution 261 plasma of
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IV. Plasma Iron Transport 263 pathw
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VI. Iron Metabolism in Cells 265 of
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VI. Iron Metabolism in Cells 267 in
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VII. Tests for Evaluating Iron Meta
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VII. Tests for Evaluating Iron Meta
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VIII. Disorders of Iron Metabolism
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References 279 ( Norrdin et al ., 2
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References 281 Fresco , R. ( 1981 )
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288 Chapter | 10 Hemostasis TABLE 1
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292 Chapter | 10 Hemostasis The pro
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294 Chapter | 10 Hemostasis exhibit
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298 Chapter | 10 Hemostasis that is
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302 Chapter | 10 Hemostasis However
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304 Chapter | 10 Hemostasis 2. Comp
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306 Chapter | 10 Hemostasis is a re
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308 Chapter | 10 Hemostasis 2. Asse
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310 Chapter | 10 Hemostasis necessi
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312 Chapter | 10 Hemostasis disease
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314 Chapter | 10 Hemostasis concent
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316 Chapter | 10 Hemostasis the rol
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318 Chapter | 10 Hemostasis proport
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320 Chapter | 10 Hemostasis a consi
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322 Chapter | 10 Hemostasis Bakhtia
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324 Chapter | 10 Hemostasis Dowd ,
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326 Chapter | 10 Hemostasis Kier ,
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328 Chapter | 10 Hemostasis Nielsen
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330 Chapter | 10 Hemostasis Tablin
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332 Chapter | 11 Neutrophil Functio
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352 Chapter | 12 Diagnostic Enzymol
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380 Chapter | 13 Hepatic Function L
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382 Chapter | 13 Hepatic Function e
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384 Chapter | 13 Hepatic Function p
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386 Chapter | 13 Hepatic Function m
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388 Chapter | 13 Hepatic Function s
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390 Chapter | 13 Hepatic Function f
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392 Chapter | 13 Hepatic Function T
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394 Chapter | 13 Hepatic Function 2
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396 Chapter | 13 Hepatic Function u
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398 Chapter | 13 Hepatic Function 2
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400 Chapter | 13 Hepatic Function c
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402 Chapter | 13 Hepatic Function F
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404 Chapter | 13 Hepatic Function A
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406 Chapter | 13 Hepatic Function E
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408 Chapter | 13 Hepatic Function K
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410 Chapter | 13 Hepatic Function R
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412 Chapter | 13 Hepatic Function W
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414 Chapter | 14 Gastrointestinal F
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Chapter 15 Skeletal Muscle Function
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II. Specialization of the Sarcolemm
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II. Specialization of the Sarcolemm
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III. Heterogeneity of Skeletal Musc
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III. Heterogeneity of Skeletal Musc
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V. Exercise and Adaptations to Trai
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VI. Diagnostic Laboratory Methods f
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VI. Diagnostic Laboratory Methods f
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IX. Selected Neuromuscular Disorder
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IX. Selected Neuromuscular Disorder
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References 479 and, recently, there
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References 481 Engel , A. G. ( 2004
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Chapter 16 Kidney Function and Dama
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II. Kidney Morphology and Function
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II. Kidney Morphology and Function
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III. Tests of Kidney Function 491 (
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III. Tests of Kidney Function 493 T
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III. Tests of Kidney Function 495 B
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6 5 4 3 2 1 0 Dog Cat Equine Cattle
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III. Tests of Kidney Function 499 d
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IV. Tests of Kidney Damage 501 1000
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IV. Tests of Kidney Damage 503 and
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IV. Tests of Kidney Damage 505 Enzy
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V. Biochemical Changes in Kidney Di
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612 Chapter | 19 Adrenocortical Fun
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624 Chapter | 20 Thyroid Function a
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626 Chapter | 20 Thyroid Function t
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628 Chapter | 20 Thyroid Function A
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630 Chapter | 20 Thyroid Function S
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634 Chapter | 20 Thyroid Function O
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636 Chapter | 21 Clinical Reproduct
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664 Chapter | 22 Trace Minerals the
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666 Chapter | 22 Trace Minerals the
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668 Chapter | 22 Trace Minerals P i
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670 Chapter | 22 Trace Minerals be
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Copper Cuproreductase Cytochromes C
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674 Chapter | 22 Trace Minerals 2 .
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676 Chapter | 22 Trace Minerals Cor
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682 Chapter | 22 Trace Minerals VI
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684 Chapter | 22 Trace Minerals red
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686 Chapter | 22 Trace Minerals of
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688 Chapter | 22 Trace Minerals Per
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Chapter 23 Vitamins Robert B. Rucke
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III. Fat-Soluble Vitamins 697 TABLE
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III. Fat-Soluble Vitamins 699 Carot
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III. Fat-Soluble Vitamins 701 Major
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III. Fat-Soluble Vitamins 703 doses
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III. Fat-Soluble Vitamins 705 Diet
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III. Fat-Soluble Vitamins 707 conta
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III. Fat-Soluble Vitamins 709 immun
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IV. Water-Soluble Vitamins 711 are
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IV. Water-Soluble Vitamins 713 abil
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IV. Water-Soluble Vitamins 715 5’
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IV. Water-Soluble Vitamins 717 H 2
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IV. Water-Soluble Vitamins 719 Enzy
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722 Chapter | 23 Vitamins When biot
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724 Chapter | 23 Vitamins Cyanocoba
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728 Chapter | 23 Vitamins nature, r
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732 Chapter | 24 Lysosomal Storage
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Chapter 25 Tumor Markers Michael D.
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II. Serum Tumor Markers 753 also be
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III. Flow Cytometry 755 cancer of t
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V. Immunohistochemistry/Immunocytoc
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V. Immunohistochemistry/Immunocytoc
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References 761 analysis will facili
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References 763 Fernandez , N. J. ,
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References 765 Meinecke , B. , and
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References 767 Walter , J. ( 2000 )
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770 Chapter | 26 Cerebrospinal Flui
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TABLE 26-7 Continued Constituent Ro
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Chapter 27 Clinical Biochemistry in
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II. Hepatotoxicity 823 Therefore, A
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III. Nephrotoxicity 825 suggested a
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IV. Toxins Affecting Skeletal and C
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VII. Toxins Affecting Erythrocytes
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VIII. Toxins Affecting Hemoglobin a
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IX. Toxins Affecting the Nervous Sy
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References 835 Plants classified as
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References 837 Solaiman , S. G. , M
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840 Chapter | 28 Avian Clinical Bio
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Budgerigar ALAT (IU/g tissue) Budge
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874 Appendix | I SI Units TABLE E S
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Appendix II Conversion Factors of S
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Appendix IV Stability of Serum Enzy
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Appendix VI Nomogram for Computing
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t0100 Appendix VIII Blood Analyte R
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884 Appendix | VIII Blood Analyte R
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890 Appendix | IX Blood Analyte Ref
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Appendix X Blood Analyte Reference
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Appendix XI Blood Analyte Reference
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Appendix XII Urine Analyte Referenc
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902 Appendix | XIII Cerebrospinal F
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904 Appendix | XIV Cerebrospinal Fl
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