Preliminary Pages

1HUMAN PHYSIOLOGY2

3HUMAN PHYSIOLOGY

Fully Revised 4th Edition

NM Muthayya MBBS MSc PhD Formerly Professor of Physiology Madurai Medical College Thanjavur Medical College Coimbatore Medical College Rajah Muthiah Medical College Annamalai University University of Health Sciences Antigua, West Indies Dean Thanjavur Medical College Dean and HOD Physiology Vinayaka Missions Kirupananda Variar Medical College Salem, India

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Human Physiology

All rights reserved. No part of this publication should be reproduced, stored in a retrieval system, or transmitted in any form or by any means: electronic, mechanical, photocopying, recording, or otherwise, without the prior written permission of the author and the publisher.

Third Edition: 2002

Revised Reprint: 2003

Fourth Edition: 2009

9788184487367

Typeset at JPBMP typesetting unit

Printed at

5Dedicated

My Students6

Foreword

This book on Human Physiology written and edited by Prof NM Muthayya, an eminent Professor of Physiology, who has contributed to the field of Physiology as the Faculty lead in various Universities in India and abroad. A revision of this textbook became inevitable for two reasons. The author has also tried to prune to some of the old wood. Even though the book has been revised with many editions, the bulk has been reduced. This edition of the book is a student friendly and easy to understand. The schematic pictures are too good. Abbreviations and symbol used in this book may be of great help to the students. Although the main framework of the textbook has been retained, almost all paragraphs have been revised.

The present edition provides additional knowledge in many sections.

The information provided in each section is exam-oriented for the benefit of the students.

Several illustrative diagrams and tables have been included to make the subject comprehensive and to revise it more easily.

Comparing to the previous edition this 4th edition is updated with reasonable additions and deletions.

The publication of the Textbook Human Physiology for medical students by Dr NM Muthayya is a timely and laudable venture. Although many costly textbooks written by western authors are available in the market, there are only very few standard textbooks prepared by Indian authors. I am glad to note that the present textbook keeps a balance between the basic essentials and advanced areas of knowledge in Physiology. I am pleased to introduce the book to the medical students and attractive to the teachers.

I have no doubt that this will be a valuable addition to the armamentarium of students of Physiology who are preparing for examination and are seeking a strong foundation. This book of Human Physiology 4th Edition will be kept for reference in the Tamil Nadu Dr MGR Medical University for the benefit of the undergraduate and postgraduate students.

As the Vice-Chancellor of the Tamil Nadu Dr MGR Medical University, I wish and congratulate Prof Dr NM Muthayya for taking much effort to bring this 4th edition of this book successfully.

Dr K Meer Mustafa Hussain8

Foreword

It gives me great pleasure to write a foreword for this comprehensive book of Physiology which incorporates latest information on Physiology.

The book “Human Physiology” is well written and tastefuly illustrated.

The author, Professor NM Muthayya, an eminent physiologist has made it possible for any beginner undergoing Medical as well as Paramedical Courses to grasp the fundamentals of Physiology through this easily readable book.

I wish to congratulate Dr Muthayya on this achievement and wish him all success.

Dr P Vijayalakshmi10

11Preface to the Fourth Edition

Since this book was published in 1986, it has undergone several modifications in its format and content. Two new editions and number of reprints have been published. This new fourth edition has been revised very extensively, new materials have been added, errors have been corrected, suggestions and information from physiology colleagues and readers have been taken into account and incorporated. Recent concepts have been included and the materials which are no longer relevant have been deleted. Number of new diagrams, tables and flow charts have been added. The format of the book is completely changed. By these changes this edition is made up-to-date and accurate to the extent possible and published under the title Human Physiology.

Inorder to emphasize the clinical significance of physiology to medical students the necessary clinical aspects have been included in relevant places then and there in the body of the subject matter and also at the end of each chapter.

To make the clinical aspects more striking and attractive these clinical aspects have been printed in italic type with red colour. The very purpose of this effort is only to initiate and induct the preclinical students to clinical studies and not to teach the clinical medicine in details which you will get during the clinical years of study.

As in the previous editions of this book, in this edition also the historical aspects, the names of Pioneer Scientists who contributed to some new discoveries and the names of Nobel Laurettes in Medicine and Physiology have been given in the respective areas in order to honour them. With an intention of creating some interest in “The History of Medicine” among students these facts have been highlighted by printing them in “Green” colour.

I am always grateful to Dr (Mrs) S Parvathi Devi, Director and Emeritus Scientist (Retd) Institute of Physiology, Madurai Medical College, Madurai, India for the encouragement she has given as my guide, friend and philosopher. I am very much thankful to Dr N Harihara Subramanian, Professor of Physiology (Retd) and Mr PV Masilamani, Artist (Retd) Madurai Medical College for their contribution of some diagrams to the first edition of this book which forms and continues as the foundation for the present new edition.

I thank profusely Mr S Pusparaj and Mrs S Maithili Yogaraj of Mythe Creators, Coimbatore for their sincere effort to make the computer designing more attractive and colourful in the present form.

I am also thankful to my family members particularly my son Engineer RM Pratheep Pratap for his help in bringing out this edition.

I thank and appreciate Shri Jitendar P Vij (Chairman and Managing Director), Mr Tarun Duneja (Director-Publishing) of Jaypee Brothers Medical Publishers (P) Ltd, New Delhi and Chennai Branch, India who brought out this book at a very short notice of time. My heart felt desire is to dedicate this book to my students from whom I got the feedback and encouragement to venture on this task.

Madurai

Tamil Nadu, India

NM Muthayya

21Abbreviations and Symbols Used in this Book µ

: Micro, 10⁻⁶

A (A)

: Angstrom unit(s) (10⁻¹⁰m, 0.1nm);also alanine

A₁, A₂, A_1B, A_2B, B, O; Major blood groups

ABC

: ATP-binding cassette

ABP

: Androgen-binding protein

ACE

: Angiotensin-converting enzyme

Acetyl-CoA

: Acetyl-coenzyme A

Ach

: Acetylcholine

ACTH

: Adrenocorticotrophic hormone

Acyl-CoA

: General symbol for an organic compound coenzyme A ester

ADH

: Antidiuretic hormone (vasopressin)

ADP

: Adenosine diphosphate

AGEs

: Advanced glycosylation end products

AHG

: Antihemophilic globulin

Ala

: Alanine

ALS

: Amyotrophic lateral sclerosis

AMP

: Adenosine 5′-monophosphate

ANP

: Atrial natriuretic peptide

APC

: Activated protein C; also antigen-presenting cell

APUD cells

: Amine precursor uptake and decarboxylation cells that secrete hormones

Arg

: Arginine

Asp

: Aspartic acid

ATP

: Adenosine triphosphate

A-V difference

: Arteriovenous concentration difference of any given substance

AV node

: Atrioventricular node

aVR, aVF, aVL

: Augmented unipolar electrocardiographic leads

AV valves

: Atrioventricular valves of heart

BGP

: Bone G1a protein

BMR

: Basal metabolic rate

BNP

: Brain natriuretic peptide

cal

: The calorie (gram calorie)

Cal

: 1000 calories; kilocalorie

cAMP

: Cyclic adenosine 3′,5′-monophosphate

CBF

: Cerebral blood flow

CBG

: Corticosteroid-binding globulin, transcortin

: Cubic centimeters

CCK, CCK-PZ

: Cholecystokinin-pancreozymin

CFF

: Critical fusion frequency

cGMP

: Cyclic 3′,5′-guanosine monophosphate

CNS

: Central nervous system

CoA

: Coenzyme A

COHb

: Carbonmonaxy hemoglobin

Compound A

: 11-Dehydrocorticosterone

Compound B

: Corticosterone

Compound E

: Cortisone

Compound F

: Cortisol

Compound S

: 11-Deoxycortisol

COMT

: Catechol-O-methyltransferase

cps

: Cycles per second, hertz

: Conditioned reflex

: Creatinine

CRH, CRF

: Corticotropin-releasing hormone

CRO

: Cathode-ray oscilloscope

: Conditioned stimulus

CSF

: Cerebrospinal fluid; also colony-stimulating factor

: Computed tomography

C terminal

: COOH end of a peptide or protein

cyclic AMP

: Cyclic adenosine 3′,5′,-monophosphate

Cys

: Cysterine

CZL

: Crystalline zinc insulin

DAG

: Diacylglycerol

: Decibel

DEA, DHEA, DHA

: Dehydroeplandrosterone22

DFP

: Diisopropyl fluorophosphate

DHT

: Dihydrotestosterone

DIT

: Dilodotyrosine

DNA

: Deoxyribonucleic acid

D₂O

: Deuterium oxide (heavy water)

DOCA

: Deoxycorticosterone acetate

DPG, 2,3-DPG

: 2,3-Diphosphoglycerate

ECF

: Extracelluar fluid

ECG, EKG

: Electrocardiogram

EDRF

: Endothelium-derived relaxing factor

EDTA

: Ethylenediaminetetraacetic acid

EMG

: Electromyogram

EPSP

: Excitatory postsynaptic potential

ERG

: Electroretinogram

FAD

: Flavin adenine dinucleotide

FEV₁

: Forced expiratory volume in first second of forced expiration after maximum inspiration

FFA

: Unesterified free fatty acid (also called NEFA, UFA)

FGF

: Fibroblast growth factor

FGFR

: Fibroblast growth factor receptor

FMN

: Flavin mononucleotide

FRH, FSH-RH, FRF

: FSH releasing hormone

FSH

: Follicle-stimulating hormone

: Foot or feet

g, gm

: Gram(s)

: Unit of force; 1 g equals the force of gravity on the earth's surface

GABA

: Gamma-aminobutyrate

GAD

: Glutamate decarboxylase

GBG

: Gonadal steroid-binding globulin

G-CSF

: Granulocyte colony-stimulating factor

GFR

: Glomerular filtration rate

: Growth hormone

GIH, GIF

: Growth hormone-inhibiting

GIP

: Gastric inhibitory peptide

Gla

: Gamma-carboxyglutamic acid

Glu

: Glutamic acid

GLUT

: Glucose transporter

Gly

: Glycine

GM-CSF

: Granulocyte-macrophage colonystimulating factor

GnRH

: Gonadotropin-releasing hormone; same as LHRH

GTP

: Guanosine triphosphate

: Hour(s)

: Histidine

: Deoxygenated hemoglobin

HBE

: His bundle electrogram

HbO₂

: Oxyhemoglobin

HCC, 25-HCC

: 25-Hydroxycholecalciferol, a metabolite of vitamin D₃

hCG

: Human chorionic gonadotropin

hCS

: Human chorionic Somatomammotropin

Hct

: Hematocrit

HDL

: High-density lipoprotein

hGH

: Human growth hormone

5-HIAA

: 5-Hydroxyindoleacetic acid

HIOMT

: Hydroxyindole-O-methyltransferase

His

: Histidine

HIV

: Human immunodeficiency virus

HLA

: Human leukocyte antigen

H substance

: Histaminelike capillary vasodilator

5-HT

: Serotonin

IDDM

: Insulin-dependent diabetes mellitus

IDL

: Intermediate-density lipoprotein

IFN

: Interferon

IGF-I, IGF-II

: Insulinlike growth factors I and II23

¹²³I-IMP

: ¹²³I-labeled iodoamphetamine

: Interleukin

lle, lleu

: Isoleucine

IP₃

: Inositol 1,4,5-triphosphate, inositol triphosphate

IPSP

: Inhibitory postsynaptic potential

: International unit(s)

IUD

: Intrauterine device

JG cells

: Juxtaglomerular cells

: Kilo-10³; see Table 3, above

kcal (Cal)

: Kilocalorie (1000 calories)

LATS

: Long-acting thyroid stimulator

LDH

: Lactate dehydrogenase

LDL

: Low-density lipoprotein

LES

: Lower esophageal sphincter

Leu

: Leucine

log

: Logarithm to base 10

LRH, LHRH, LRF

: Luteinizing hormone-releasing hormone; same as GnRH

LTD

: Long-term depression

LTP

: Long-term potentiation

: Molarity (mol/L); also mega-, 10⁻⁶; also methionine

M cells

: Microfold cells

MAO

: Monoamine oxidase

MBC

: Maximal breathing capacity (same as MVV)

M-CSF

: Macrophage colony-stimulating factor

MDMA

: 3,4-Methylenedioxy methamphetamine

MHC

: Major histocompatibility complex; also myosin heavy chain

MHPG

: 3-Methoxy-4-hydroxyphenylglycol

MRI

: Magnetic resonance imaging

mRNA

: Messenger RNA

MSH

: Melanocyte-stimulating hormone

MVV

: Maximal voluntary ventilation

: Normality (of a solution): also Newton (SI unit of force); also asparagine

NAD

: Nicotinamide adenine dinucleotide; same as DPN

NADH

: Dihydronicotinamide adenine dinucleotide; same as DPNH

NADP+

: Nicotinamide adenine dinucleotide phosphate; same as TPN

NADPH

: Dihydronicotinamide adenine dinucleotide phosphate; same as TPNH

NGF

: Nerve growth factor

NIDDM

: Non-insulin-dependent diabetes mellitus

NMDA

: N-Methyl-<sc>D</sc>-aspartate

: Nitric oxide

NREM sleep

: Nonrapid eye movement (spindle) sleep

NSAID

: Nonsteroidal anti-inflammatory drug

NTS

: Nucleus of the tractus solitarius

OVLT

: Organum vasculosum of the lamina terminalis

: Pico-, 10⁻¹²;

P450

: Cytochrome P450

P₅₀

: Partial pressure of O₂ at which hemoglobin is half-saturated with O₂

PAF

: Platelet-activating factor

PAH

: Para-Aminohippuric acid

PBI

: Protein-bound-iodine

P cells

: Principal cells in the renal tubules; also pacemaker cells of SA and AV nodes

PCD

: Programmed cell death, apoptosis

PDECGF

: Platelet-derived endothelial cells growth factor

PET

: Positron emission tomography

PGO spikes

: Ponto-geniculo-occipital spikes in REM sleep24

: Negative logarithm of the H⁺ concentration of a solution

PIH, PIF

: Prolactin-inhibiting hormone

: Negative logarithm of the equilibrium constant for a chemical reaction

PLC

: Phospholipase C

PRH, PRF

: Prolactin-releasing hormone

PTA

: Plasma thromboplastin antecedent (clotting factor XI)

PTC

: Plasma thromboplastin component (clotting factor IX); also phenylthiocarbamide

PTH

: Parathyroid hormone

PZI

: Protamine zinc insulin

RAS

: Reticular activating system

rbc

: Red blood cell(s)

RDS

: Respiratory distress syndrome

REM sleep

: Rapid eye movement (paradoxical) sleep

RNA

: Ribonucleic acid

RPF

: Renal plasma flow

: Respiratory quotient

: second(s); also standard deviation of a sample

SA node

: Short-chain fatty acid(s)

SCUBA

: Self-contained underwater breathing apparatus

SDA

: Specific dynamic action

SGLT 1

: Sodium-dependent glucose transporter 1

SGOT

: Serum glutamic-oxaloacetic transamines

SIDS

: Sudden infant death syndrome

SIF cells

: Small, intensely fluorescent cells in sympathetic ganglia

SPCA

: Proconvertin (clotting factor VII)

SS 14

: Somatostain 14

SS 28

: Somatostatin 28

SS 28 (1–12)

: Polypeptide related to somatostatin that is found in tissues

STH

: Somatotropin, growth hormone

T₃

: 3,5,3,-Triiodothyronine

T₄

: Thyroxine

TBG

: Thyroxine-binding globulin

TBPA

: Thyroxine-binding prealbumin (now called transthyretin)

TBW

: Total body water

Tc cells

: Cytotoxic T cells

TEA

: Tetraethylammonium

TETRAC

: Tetraiodothyroacetic acid

TF/P

: Concentration of a substance in renal tubular fluid divided by its concentration in plasma

TGF

: Transforming growth factor

: Renal tubular maximum

TNF

: Tumor necrosis factor

TSH

: Thyroid-stimulating hormone

T/s ratio

: Thyroid/serum iodide ratio

TTX

: Tetrodotoxin

Tyr

: Tyrosine

: Unit(s)

UFA

: Unesterified free fatty acid (same as FFA)

: Unconditioned stimulus

UTP

: Uridine triphosphate

V₁, V₂, etc

: Unipolar chest electrocardiographic leads

VIP

: Vasoactive intestinal polypeptide

VLDL

: Very low density lipoprotein

VMA

: Vanillymandelic acid (3-methoxy-4-hydroxy-mandelic acid)

VOR

: Vestibulo-ocular reflex

: Unipolar right arm electrocardiographic lead

wbc

: White blood cell(s)

X chromosome

: One of the sex cortex in humans

Y chromosome

: One of the sex chromosomes in humans25

Greek Alphabet
α	alpha	ν	nu
β	beta	ξ	xi
γ	gamma	ο	omicron
δ	delta	π	pi
ε	epsilon	ρ	rho
ζ	zeta	σ, ς	sigma
η	eta	τ	tau
θ	theta	υ	upsilon
ι	iota	φ	phi
κ	kappa	χ	chi
λ	lambda	ψ	psi
μ	mu	ω	omega

Chapter Notes