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th13	Edition
Revised	&	Updated
Organic
Evolution
(Evolutionary	Biology)
Veer	Bala	Rastogi
MEDTECH
A	Division	of
Scientific	International
Evolution
(Evolutionary	Biofogy)
THIRTEENTH	EDITION
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•	Plant	Growth	and	Development	.	Anwa	r	Hussain
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•	Food	Microbiology	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
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.	.	.	.	.	.	.	.	.	.	.	.	.	Michael	d.	Lagrega,	Phillip	Monika	Rustagi	R.S.	Singh	.	.	.	R.S.	Singh	...	R.S.	Singh	Veer
Bala	Rastogi
Evolution
(Evolutionary	Biology)
THIRTEENTH	EDITION
Dr.	Veer	Bala	Rastogi
M	.Sc.	(Gold	Medalist)	Ph.D.,	F.A.Z.	Formerly	Reader,	Department	of	Zoology
Gargi	College,	University	of	Delhi,	Delhi
Ex-Member,
Textbook	Evaluat	ion	Committee
NCERT,	New	Delhi
Recipient	ofDistinguished	Author	Award	2012
by	the	Federation	ofEducation	publishers
in	India,	Delhi
MEDTECH
A	Division	of	Scientific	International
Engaging	Sciences-Developing	Mindsl
Evolution
(Evolutionary	Biology)
MEDTECH
ADivision	of
Scientific	International	Pvt.	Ltd.
Copyright	©	Author
Thirteenth	Edition:	2018
All	rights	reserved.	No	Part	of	this	publication	may	be	reproduced	or	transmitted
in	any	form	or	by	any	means–electronic	or	mechanical,	including	photocopy,
recording,	or	any	information	storage	and	retrieval	system–without	permission	in
writing	from	the	publisher.
Disclaimer:	Every	effort	has	been	made	to	avoid	any	error	or	omission	in	this
publication.	It	may	be	noted	that	neither	the	author	nor	the	publisher	will	be
responsible	for	any	damage	or	loss	of	action	to	any	one	of	any	kind,	in	any
manner,	therefrom.
ISBN:	978-93-87465-32-9
The	authors,	editors,	contributors	and	the	publisher	have,	as	far	as	it	is	possible,
taken	care	to	ensure	that	the	information	given	in	this	text	is	accurate	and	up-to-
date.	However,	readers	are	strongly	advised	to	confirm	that	the	information
complies	with	current	standards	of	practice.
Every	effort	has	been	made	where	necessary	to	contact	holders	of	copyright	to
obtain	permission	to	reproduce	copyright	material.	If	any	have	been
inadvertently	overlooked,	the	publisher	will	be	pleased	to	make	the	necessary
arrangements	at	the	first	opportunity.
Published	by:	Vinod	Kumar	Jain,	Scientific	International	(Pvt.)	Ltd.
Registered	Office:	Branch	Offices:
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SIPL	EDITORIAL/PRODUCTION	TEAM	Production	Manager	:Sunil	K.
Panda
Publishing	Manager	(Operations):Vinod	Chauhan	Cover	Design:Ashis	Dhar
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Printed	in	India
Preface
Evolutionary	Biology	is	as	wide	as	the	world	of	animals	and	plants,	past	and
present.	Recent	years	have	seen	an	immense	expansion	of	evolutionary	studies	to
many	areas	that	were	scarcely	touched	before.	The	last	twenty	five	years	have
witnessed	the	revolutionary	impact	of	molecular	biology	upon	genetics	and
developmental	biology,	the	two	fields	of	fundamental	importance	for
evolutionary	studies.	As	a	result	evolutionists	have	acquired	new	tools	and
concepts	for	investigating	evolutionary	processes.	Due	to	rapid	progress	made	in
the	field	of	molecular	biology,	many	unexpected	and	revealing	discoveries	have
already	emerged.	Further	molecular	studies	will	certainly	modify	the	paradigm
of	evolution	as	we	now	understand	it	although	the	extent	of	change	remains	to	be
determined.
So	,	why	did	I	write	a	book	about	evolution?	Because,	while	teaching	evolution,
I	observed	that	a	lot	of	young	students	are	confused	about	exactly	what	evolution
is,	what	it	does	,	how	it	works,	and	why	it	is	important.	This	book	will	help	all
such	readers	to	everything	out.	The	content	of	the	book	is	meant	to	reflect	a	wide
range	of	evolutionary	principles	and	to	offer	examples	showing	how
evolutionary	forces	work	and	have	worked	in	past	to	give	us	such	a	colourful
and	varied	world.	The	text	is	intended	to	help	the	students	develop	a	large
number	of	ideas	or	centres	of	interest	of	his	own	concerning	evolution.
The	text	in	this	book	is	divided	into	five	parts.	Part	I,	begins	with	the	Concept	of
Evolution,	which	deals	with	the	meaning	of	word	'Evolution',	and	the	historical
development.	It	provides	a	comprehensive	account	of	evidences	to	support
concept	of	evolution,	and	different	theories	for	exploring	the	mechanism	of
evolution.	Part	II	deals	with	the	mechanism	of	evolution.	Topics	on	Variation
and	Gene	Mutations	which	form	the	necessary	substrate	for	natural	selection	are
dealt	in	a	simple	and	understandable	form.	Next	two	chapters	deal	with
chromosomal	changes.	Chromosomal	aberrations	include	changes	in	the
arrangement	of	genes	in	a	chromosome	and	variations	in	the	chromosome
number.	Both	types	of	chromosomal	variations	advocate	that	no	two	organisms
or	parts	of	organism	are	precisely	alike	because	variations	influence	the
characters	of	individuals.	Next	chapter	is	on	Isolation	which
vi	~	Preface
deals	with	isolating	mechanisms	that	split	the	species	populations	into	separate
groups	and	their	evolution	into	distinct	species.
Part	III	includes	chapters	on	Population	,	Population	Genetics	and	Persistence	of
Variability	within	populations	leading	to	the	formation	of	species.	It	is	followed
by	Genetic	drift	and	gene	flow	among	populations.	The	chapter	on	Natural
selection	in	action	stresses	that	the	evolutionary	agents	bring	in	changes	in	the
frequencies	of	alleles	and	genotypes	in	Mendelian	populations	that	produce	more
efficient	adaptive	relationship	with	the	environment	to	ensure	better	survival	and
comparative	reproductive	success	.	Next	chapter	deals	with	evolution	of	genes
and	genomes.	This	chapter	discusses	how	the	evolutionists	with	the
advancement	in	the	knowledge	of	molecular	genetics	are	able	to	study
evolutionary	changes	in	the	genes,	genomes	and	gene	pools	of	populations	and
how	the	organisms	undergo	mutations	and	get	adapted	to	new	environments.
Part	IV	emphasises	on	the	basic	patterns	of	evolution	.	The	chapter	on
Microevolution	and	Macroevolution	throw	s	light	on	the	interaction	of	the
elemental	forces	of	evolution,	i.e.,	mutations,	variation,	natural	selection,	genetic
drifts	leading	to	microevolution.	Microevolution	explains	the	origin	of	new
adaptive	types	through	a	process	of	population	fragmentation	.	Next	chapter	is
on	Patterns	of	Evolution,	i.e.,	sequential	and	divergent	evolution,	phyletic
gradualism	and	punctuated	equilibrium,	anagenesis	and	cladogenesis;
monophyletic,	polyphyletic	and	paraphyletic	evolution,	divergent	evolution,	and
conditions	responsible	for	adaptive	radiation	and	the	causes	and	significance.
This	is	followed	by	convergent	evolution	or	adaptive	convergence	supplemented
with	examples	of	adaptive	convergence.	It	is	followed	by	coevolution	which	is
also	a	pattern	of	evolution	in	which	two	interacting	species	influence	each	other's
adaptive	changes	over	the	time,	its	examples	and	outcome	of	coevolution.
Chapter	on	adaptations	explains	how	the	organisms	enable	themselves	to	thrive
successfully	in	a	particular	environment.
Part	V	provides	an	insight	into	the	origin	and	history	of	life	on	Earth.	Chapter	on
'Origin	of	Life'	covers	the	physical	and	chemical	environment	that	existed	on
ancient	primitive	Earth	and	supported	the	origin	of	first	living	form,	the
beginning	of	biological	evolution	and	gradual	complexity	attained	there	in.
Chapter	on	fossil	records	reveals	the	long	history	of	life	which	had	been	partly
recorded	in	certain	rocks	of	earth's	crust	in	the	form	of	fossils.	Next	chapter
summarises	the	whole	evolutionary	story	as	shown	by	fossils	in	geological
records.	Last	chapter,	Evolution	of	Primates	and	Man	is	written	from	zoological
point	of	view,	for	the	student	needs	to	know	first	something	about	what	actually
happened.
I	have	been	extra	cautious	in	presenting	the	text	in	easy	and	lucid	language.
Diagrams	are	specially	designed	and	also	taken	from	various	sources	for	clarity
and	simplicity.
I	am	pleased	to	record	thanks	to	universities	all	over	the	country	who	my
colleagues	at	various	institutions	and	have	provided	immense	valued	help	while
preparing	this	endeavour.	I	request	all	of	them	to	send	their	valued	suggestions
and	critical	review	in	the	subsequent	edition.
Preface	~	vii
I	am	highly	thankful	to	Sh.	Rajan	Jain	(Director)	Scientific	International	Pvt.
Ltd.,	Sh.	Vinod	Chauhan	(HOD-Production)	and	Sh.	S.K.	Panda	(Editor-
cumCoordinator)	for	their	tiredless	support	throughout	in	giving	the	shape	to	the
book	what	it	is	with	you.
Veer	Bala	Rastogi
Contents
Prefa	ce	v
UNIT	I.	CONCEPT,	EVIDENCES	AND	THEORIES	OF	EVOLUTION	1	1.
Concept	of	Evolution	3
1.1	Evolution	and	Evolutionary	Biology	3
1.2	Definition	of	Biological	Evolution	5
1.3	Basic	Concept	of	Organic	Evolution	5
1.4	Development	of	the	Idea	of	Evolution	6
1.5	Evolution-A	Fact	or	Just	a	Theory	14
Ke	y	Terms	15
Review	Questions	16
Furth	er	Readings	16
2.	Evidences	for	Evolution	17
2.1	Doctrine	of	Biological	Evolution	17
2.2	Evidences	for	Biologica	l	Evolution	17
2.3	Evidences	from	Comparative	Anatomy	and	Morphology	(Tectology)	18	2.4
Evidences	from	Vestigial	Organs	28
2.5	Evidences	from	Atavism	or	Reversion	30
2.6	Evidences	from	Comparative	Embryo	logy	32
2.7	Evidences	from	Palaeonto	logy	or	Palaeobiology	38
2.8	Evidences	from	Geographical	Distribution	or	Biogeographical
Evidences	51
2.9	Evidences	from	Connecting	Links	55
2.10	Evidences	from	Taxonomy	58
2.11	Evidences	from	Biochemistry	and	Physiology	60
x	~	Contents
2.12	Evidences	from	Molecular	Records	62
2.13	Evidences	from	Cytology	64
2.14	Evidences	from	Gen	etics	66
Key	Terms	67
Review	Questions	68
Further	Readings	69
3.	Theories	of	Evolution	70
3.1	Lamarck	and	Lamarckism	70
3.2	Inheritance	of	Acquired	Characters	(Lamarckism)	70
3.3	Darwinism	or	Theo	ry	of	Natu	ral	Selection	76
3.4	Mutation	Theory	of	Evolution	92
3.5	Modem	Synthetic	Theory	of	Evolution	or	The	Evolutionary	Synthesis	95
3.6	Neutral	Theory	of	Evoluti	on	99
Key	Terms	100
Review	Questions	100
Further	Readings	101
UNIT	II.	MECHANISMS	OF	EVOLUTION	103
4.	Variation	105
4.1	Variation	and	Variability	105
4.2	Nature	of	Variation	105
4.3	Types	of	Variation	105
4.4	Sources	of	Variation	108
4.5	Variation	in	Number	of	Chromosomes	(Heteroploidy)	109
4.6	Chromosomal	Aberrations	11	2
4.7	Gen	e	Mutations	or	Point	Mutations	11	4
4.8	Mendelian	Recombination	or	Sexual	Recombination	116
4.9	Recombination	due	to	Exchange	of	Genes	between	Homologous
Chromosomes	of	a	Pair	11	8
4.10	Hybridi	sati	on	120
Key	Terms	122
Review	Questions	122
Further	Readings	122
5.	Gene	Mutations	124
5.1	Definition	124
5.2	History	124
5	.3	Characteristics	of	Mutations	125
Contents	~	xi
5.4	Kinds	of	Mutations	126
5.5	Causes	of	Mutations	or	Mutagenic	Agents	129
5.6	Molecu	lar	Basis	of	Single	Gene	Mutations	or	Point	Mutations	131
5.7	Substitution	Mutations	133
5.8	Frameshift	Mutations	142
5.9	Mutation	Rates	144
5.10	Effects	of	Mutations	on	Fitness	146
5.	11	Randomness	of	Mutations	147
5.12	Mutations	and	Genetic	Polymorphism	148
5.13	Mutations	and	Evolution	148
Key	Terms	149
Review	Questions	149
Furth	er	Readings	150
6.	Chromosomal	Aberrations	151
6.1	Chromosomal	Aberrations	lS	I
6.2	Origin	of	Chromo	soma	l	Aberration	s	151
6.3	Types	of	Chromosomal	Aberrations	lSI
6.4	Deletion	or	Deficiency	152
6.5	Duplication	or	Repeat	156
6.6	Inversion	s	160
6.7	Translocation	166
Key	Terms	172
Review	Questions	173
Furth	er	Readings	174
7.	Variation	in	Chromosome	Number	(Heteroploidy)	175
7.1	Changes	Involving	Entire	Set	of	Chromosomes	(Euploidy)	175
7.2	Changes	Involving	Number	of	Chromosomes	in	a	Set	(Aneuploid	y)	176
7.3	Euploidy	176
7.4	Origin	of	New	Species	Through	Polyploidy	or	Evolutionary	Role	of
Polyploidy	183
7.5	Induced	Polyploidy	187
7.6	Aneup	loidy	188
Key	Terms	192
Review	Questions	192
Furth	er	Readings	193
8.	Reproductive	Isolating	Barriers	194
8.1	Introduction	194
8.2	History	194
xii	~	Contents
8.3	Role	of	Reproductive	Isolation	195
8.4	Types	of	Reproductive	Isolating	Mechanisms	195
8.5	Premating	Barriers	196
8.6	Postmating	Prezygotic	Barriers	203
8.7	Postzygotic	Barriers	205
8.8	Multiple	Isolating	Barriers	207
8.9	Genetic	Basis	of	Reproducti	ve	Barriers	or	Reproductive	Isolation	208	8.10
Origin	of	Reproductive	Isolat	ion	and	Origin	of	Species	210	8.11	Evolution	of
Reproductive	Barriers	211
Key	Terms	212
Review	Questions	212
Further	Readings	213
UNIT	III.	SPECIATION	215
9.	Population	Genetics,	Gene	Frequencies
and	Hardy-Weinberg	Equilibrium	217
9.1	Population	Genetics	217
9.2	Population	217
9.3	Gene	Pool	219
9.4	Fundamental	Principles	of	Genetic	Variation	in	Populations	220	9.5	Hardy-
We	inberg	Equilibrium	226
9.6	Hardy-Weinberg	Principle	and	Evolution	(Factors	that	Change	Gene
Frequency	)	232
9.7	Genetic	Landscape	of	a	Population	and	Evolution	236
Key	Terms	238
Review	Questions	238
Further	Readings	239
10.	Persistence	of	Variability	within	Populations:	Polymorphism	241	10.1
Variability	within	Populations	241
10.2	Polymorphism	241
10.3	Balanced	Polymorphism	243
10.4	Transient	Polymorphism	249
10.5	Origin	of	Polymorphism	250
10.6	Mechanisms	to	Maintain	Polymorphism	within	Populations	250
K	ey	Terms	252
Review	Questions	252
Further	Readings	253
Contents	[i]	xiii	11.	From	Population	to	Species	(Speciation)	254
11	.1	Species	and	Speciation	254
11	.2	The	Species	Concept	254
11.3	Species	Categories	260
11.4	Origin	of	Species	(Speciation)	262
11	.5	Modes	of	Speciations	262
11.6	Allopatric	Speciation	265
11	.7	Peripatric	Speciation	of	Marginal	Populations	270
11.8	Parapatric	Speciation	271
11	.9	Alloparapatric	Speciation	272
11.10	Sympatric	Speciation	272
11	.11	Consequences	of	Speciation	277
11.12	Rate	of	Speciation	277
11.13	Factors	Responsible	for	Variation	in	Speciation	Rates	277
11.14	Theories	of	Speciation	278
Key	Terms	280
Review	Questions	280
Further	Readings	281
12.	Genetic	Drift	and	Gene	Flow	283
12.1	Random	Genetic	Drift	or	Sewall	Wright	Effect	283
12.2	Theory	of	Genetic	Drift	283
12.3	Salient	Features	of	Genetic	Drift	284
12.4	Genetic	Basis	of	Random	Genetic	Drift	286
12.5	Genetic	Drifts	in	Real	Populations	287
12.6	Founder	Effect	or	Founder	Principle	289
12.7	Bottleneck	Phenomenon	291
12.8	Genetic	Drift	and	Evolution	293
12.9	Gene	Flow	295
K	ey	Terms	297
Review	Questions	297
Further	Readings	298
13.	Natural	Selection	in	Action	299
13.1	Concept	of	Natural	Selection	299
13.2	Salient	Features	of	Natural	Selection	302
13.3	Natural	Selection	in	Nature	302
13.4	Demonstration	of	Role	of	Natural	Selection	306
13.5	Working	of	Natural	Selection	307
13.6	Components	of	Natural	S	election	or	Levels	of	Natural	Selection	308
13	.7	The	Results	of	Natural	Selection	312
13.8	Models	of	Selection	313
13.9	Frequency	Dependent	Selection	322
13.10	Heterozygous	Advantage	or	Heterosis	322
13.11	Balancing	Selection	and	Balanced	Polymorphism	323
13.12	r-Selection	and	k-Selection	324
13.13	Selection	Pressure	or	Selection	Intensity	325
13.14	Selection	and	Reproduction	326
13.15	Selection	and	Mutations	327
13.16	Selection	and	Variation	329
13.17	Selection	and	Adaptations	(The	Baldwin	Effect)	330
Key	Terms	330
Review	Questions	331
Further	Readings	331
14.	Evolution	of	Genes	and	Genomes	333
14.1	Molecular	Evolution	333
14.2	Molecular	Phylogenie	s	333
14.3	Proteins	and	Phylogenetic	Relationship	334
14.4	Origin	and	Evolution	of	New	Genes	336
14.5	Regulatory	Genes	and	Evolution	345
14.6	Nucleic	Acid	Phylogenies	345
14.7	Genome	and	Phylogenetic	Relationship	347
14.8	Convergent	Molecular	Evolution	347
14.9	Molecular	Clocks	or	Evolutionary	Clocks	348
14.10	Molecular	Evolution	in	Test	Tube	351
Key	Terms	352
Review	Questions	352
Further	Readings	353
UNIT	IV.	BASIC	PATTERNS	OF	EVOLUTION	355
15.	Patterns	of	Evolution	357
15.1	Sequential	and	Divergent	Evolution	357
15.2	Phyletic	Gradualism	and	Punctuated	Equilibrium	358
15.3	Anagenesis	and	Cladogenesis	359
15.4	Monophyletic,	Polyphyletic	and	Paraphyletic	Evolution	361
15.5	Divergent	E	volution	(Adaptive	Radiation	or	Adaptive	Divergence)	362
15.6	Convergent	Evolution	or	Adapti	ve	Convergence	or	Parallel
Evolution	369
Contents	~	XV	15.7	Coevolution	372
Key	Terms	372
Review	Questions	372
Further	Readings	373
16.	Microevolution	and	Macroevolution	374	16.1	Microevolution	374
16.2	Macroevolution	(Adaptive	Radiation)	378	16.3	Megaevolution	383
16.4	Trends	During	Macro	and	Mega-Evolution	385
Key	Terms	385
Review	Questions	385
Further	Readings	386
17.	Adaptations	387
17.1	Introduction	387
17.2	Definition	of	Adaptations	387
17.3	Kinds	of	Adaptations	388
17.4	Mimicry	395
17.5	Batesian	and	Mullerian	Mimicry	400
17.6	Co-adaptation	402
17.7	Animal	Association	Adaptations	404
17.8	Biotic	Adaptations	and	Organismic	Adaptations	404	17.9	Preadaptations
and	Postadaptations	405
17	.10	r-Adaptations	406
17.11	k-Adaptations	407
Key	Terms	407
Review	Questions	408
Further	Readings	408
UNIT	V.	FOSSILS	AND	HISTORY	OF	LIFE	ON	EARTH	411
18.	Origin	of	Life	on	Earth	413
18.1	Origin	of	Life	(Biopoiesis)	413
18.2	Ancient	and	Medieval	Beliefs	413
18	.3	Modem	Hypothesis	of	Origin	of	Life	or	Biochemical	Origin	of
Life	416
18.4	Biochemical	or	Chemosynthetic	Origin	of	Life	421
18.5	The	Earliest	Cells	436
18.6	Where	Life	Originated?	437
18.7	Earliest	Evidence	of	Existence	of	Life	on	Earth	438
18.8	Evolution	o	f	Eukaryotic	Organelles	439
18.9	Life	on	other	Planets	439
Key	Terms	443
Review	Questions	444
Further	Readings	445
19.	History	of	Life	on	Earth	446
19.1	Geological	Time	Scale	446
19.2	Azoic	Era	452
19.3	Archeozoic	Era	453
19.4	Proterozoic	Era	(The	Era	of	Former	Life)	453	19.5	Palaeozoic	Era	454
19.6	Mesozoic	Era	(Era	of	Intermed	iate	Life)	462	19.7	Coenozoic	Era	(Era	of
Recent	Life)	474
Key	Terms	478
Review	Questions	478
Further	Readings	478
20.	Fossils	and	Fossil	Records	480
20.1	Earth's	Structure	480
20.2	Classification	of	Rocks	480
20.3	Fossils	486
20.4	Exposing	Fossils	492
20.5	Interpretation	of	Fossil	Records	493
20.6	Law	of	Superposition	494
20.7	Williston's	Rule	494
20.8	Cope's	Rule	495
20.9	Allometry	(Differential	Growth	Rate)	497
20.10	Determination	of	Age	of	Fossil	s	or	Dating	of	Fossils	497
20.11	Significance	of	Study	of	Fossils	501
20.12	Incompleteness	of	Fossil	Records	503
20.13	Evolutionary	Rate	Through	Fossil	Records	505	Key	Terms	509
Review	Questions	509
21.	Origin	and	Evolution	of	Horse	510
21.1	Place	and	Time	of	Origin	510
21.2	Evolutionary	Trends	510
21.3	Characteristics	of	Modem	Horse	512
Contents	~	xvii
21.4	Phylogeny	512
21.5	Side	Lines	519
Key	Terms	520
Review	Questions	520
22.	Origin	and	Evolution	of	Man	521
22.1	Introduction	521
22.2	Scienti	sts	Associated	with	Human	Evolution	521
22.3	Time	of	Origin	of	Primates	and	Man	523
22.4	Place	of	Origin	of	Man	524
22.5	Primate	Heritage	525
22.6	Special	Features	of	Primates	525
22.7	Evolution	and	Adaptive	Radiation	in	Primates	526
22.8	Compelling	Causes	of	Evolution	of	Man	528
22.9	Impact	of	the	Descent	from	Tree	s	on	Primate	Organisation	528	22.10
Evolutionary	Trends	during	Human	Evolution	529
22.11	Evidences	from	Molecular	Biology	in	Support	of
Hominid	Evolution	from	Apes	535
22.12	Common	Ance	stor	s	of	Apes	and	Man	in	Oligocene	and	Miocene	538
22.13	Common	Ancestor	s	of	Apes	and	Man	in	Pliocene	Period	,540	22.14
Evolution	of	Man	in	Pleistocene	Period	541
22.15	Cultural	Evolution	of	Human	s	551
22.16	Impa	ct	of	Evolution	on	Human	Brain	551
22.17	Human	Races	552
22.18	Archa	eological	Division	s	of	Pleistocene	and	Holocene	Periods	553	22.19
Monophyletic	or	Polyphyletic	Origin	of	Man	554
22.20	Punctuated	Equilibrium	in	Human	Evolution	556
Key	Terms	557
Review	Questions	557
Further	Readings	558
Glossary	559
UNIT-I
Concept,	Evidences	and	Theories	of	Evolution
Chapter	1.	Concept	of	Evolution	Chapter	2.	Evidences	for	Evolution	Chapter	3.
Theories	of	Evolution
1
concept	of	Evolution
1
.1	EVOLUTION	AND	EVOLUTIONARY	BIOLOGY
1.1.1	Evolution	(L.	evolvere	=to	roll	or	to	unfold)
English	philosopher	Herbert	Spencer	(1820	-1903)	coined	the	term	'	evolution'	to
represent	the	phenomenon	that	brings	about	continuous	and	orderly	changes	in
nature	.	The	word	'evolution'	was	derived	from	Latin	word	evolvere	where	'e'
means	out	and	volvere	means	to	roll	or	unfold.
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s:	o
Leaves	on	tree	change	colour	and	fall	over	several	weeks
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£;
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Mountain	ranges	erode	over	millions	of	years
FIG.	1.1:	Difference	in	the	biological	and	physical	evolution	in	time.
E	volution	is	described	as	change	through	time.	It	can	be	used	to	represent	any
change	in	physical	or	biological	world	.	Lots	of	things	in	our	surroundings
change	over	the	time	:	the	leaves	on	trees	change	colour	and	fall,	plants	grow
and	die,	mountain	ranges	rise	and	erode	,	languages	and	cultures	change	.	As	a
matter	of	fact	evolution	occurs	at	different	levels	and	involves	all	the
components	of	universe	both	living	or	nonliving.	It	may	be:
•	at	the	molecular	level	(chemical	evolution)
•	at	the	level	of	physical	objects	like	change	in	land	topography	(physical
evolution)
•	at	the	level	of	stars	and	planets	in	the	universe	(stellar	evolution	or	cosmic
evolution)
•	at	the	level	of	living	objects	(biological	evolution)
1.1.2	Evolutionary	Biology	or	Biological	Evolution
E	volutionary	Biology	is	defined	as	the	process	of	gradual	changes	in	organisms
to	form	more	and	more	complex	forms	over	a	long	period	of	time.
•	Darwin	has	defined	evolution	as	'Descent	with	modification.'
•	Theodor	Dobzhansky	has	defined	'evolutionary	biology'	as	the	study	of
history	of	evolution	of	newer	and	more	complex	forms	of	life	on	the	Earth	from
pre-existing	simpler	ones	over	a	period	of	time.
The	term	'bioevoluti	on'	or	'evolutionary	biology'	was	introduced	by	Mayr
(1970).	It	is	also	called	organic	evolution	or	biological	evolution.
Biological	evolution	is	not	just	a	change	in	time.	It	deals	with	very	specific	type
of	changes	such	as	changes	in	the	frequency	of	different	genes	in	the	organisms
of	a	population	or	species	over	generations	or	large-scale	changes	leading	to	the
origin	of	new	species	from	a	common	ancestor	over	many	generations	.
The	central	idea	of	biological	evolution	is	that	all	life	on	Earth	shares	a	common
ancestor,	just	as	you	and	your	cousins	share	a	common	grandmother	or
grandfather.
Through	the	process	of	descent	with	modification,	the	common	ancestor	of	life
on	Earth	gave	rise	to	tremendous	diversity	over	a	veryvery	long	period.	This
diversity	in	life	is	well	illustrated	by	present	day	living	forms	and	the	forms	that
existed	in	past	and	are	documented	in	the	fossil	records.
Short-term	change
Change	through	time	•
Common	ancestor
A	genealogy	illustrates	change	with	inheritance	over	a	small	number	of	years
A	B
FIG.	1.2:	A.	Short-term	changes	are	seen	in	family	genealogy.	They	introduce
minor
differences	but	do	not	lead	to	evolution	;	B.	Long-term	changes	in	the	organisms
of	common	ancestors	,	on	accumulation	for	very	long	period	produce
tremendous	diversity	in	living	forms	and	lead	to	evolution	of	new	form	by
descent	with	modification.
Concept	of	Evolution	IiJ	5
Based	on	the	concept	of	biological	evolution,	all	living	organisms,	plants,
animals	and	bacteria,	etc.,	have	some	common	ancestor.	We	can	say	that	not	only
monkeys	or	chimpazees	even	birds,	whales,	snakes,	worms	and	trees	are	all	our
distant	cousins	.
1.2	DEFINITION	OF	BIOLOGICAL	EVOLUTION
Biological	evolution	or	organic	evolution	is	defined	as	'the	process	of	continuity
of	life	with	constant	modifications.'	It	means	living	organisms	modify	and	adapt
according	to	the	everchanging	environmental	needs.	These	modifications	keep
accumulating	in	the	organisms	generations	after	generations,	resulting	in	more
complex	and	better	adapted	new	species.	Therefore,	organic	evolution	is	the
evolution	of	present	complex	and	highly	organised	living	beings	from	simpler
and	less	organised	living	beings	of	the	past	by	gradual	modifications
accumulated	through	successive	generations	over	millions	of	years.
1.3	BASIC	CONCEPT	OF	ORGANIC	EVOLUTION
The	basic	concept	of	organic	evolution	envisages	'continuity	of	life	with	constant
modification'.	It	suggests	that:
•	Environmental	conditions	in	nature	are	everchanging.
•	Organisms	have	an	inherent	tendency	to	change	in	response	to	the	changing
environmental	conditions.	This	is	called	adaptability	or	adaptation.
•	Such	adaptive	changes	in	organisms	are	inherited	by	the	offspring	and	lead	to
the	'Origin	of	new	species'	(Evolution).
•	Since	changes	in	the	organisms	are	due	to	adaptations,	new	species	are	always
better	adapted	and	more	organised	than	their	ancestors.
•	Different	members	of	a	species,	on	being	adapted	to	different	environments,
diversify	and	evolve	along	several	divergent	lines	and	form	new	species.
•	All	the	present	day	species	had	a	common	ancestor	at	some	or	other	time	of
their	evolution	(Monophyletic	genealogy).
•	Individuals	migrate	from	their	place	of	origin	to	varied	geographical	areas	and
gradually	adapt	to	different	environmental	conditions.	This	results	in	the
formation	of	several	new	species	from	one	ancestral	species	(Divergent
evolution).
•	Organisms	from	varied	regions	also	migrate	to	a	common	habitat	and	modify
to	adapt	to	that	habitat.	As	a	result	organisms	from	distantly	related	groups
develop	common	features	(Convergent	evolution).
•	Evolution	is	a	very	complex	and	extremely	slow	process.	It	is	not	possible	to
see	one	type	of	animals	changing	to	other,	but	presence	of	integrading	organisms
supports	the	concept	of	evolution.
•	Evolutionary	changes	are	continuous.	They	occurred	in	past,	are	continuing	in
present,	and	will	continue	in	future.
1.4	DEVELOPMENT	OF	THE	IDEA	OF	EVOLUTION
Charles	Darwin's	name	is	c	losel	y	associated	with	the	concept	of	evolution	and
for	many	people	Darwinism	is	evolution,	but	the	concept	of	evolution,	for	the
first	time	appeared	in	the	writings	of	ancient	Greeks.	Their	explanation	of	the
origin	of	living	things	as	given	by	Empedocles	and	Anaximander	supported	the
notion	of	dynamic	world	and	rep	laced	the	mythological	explanation.
1.4.1	Greek	Theories
I.	Thales	(624-548	BC)	propounded	the	theory	of	aquatic	or	marine	orgin	of	life.
2	.	Anaximander	(611-547	BC)	was	call	ed	'the	earlies	t	evo	lutionist'	by	Osborn
(1894).	He	proposed	that	all	living	beings	have	arisen	from	a	primordial	fluid	or
slime	to	which	they	ultimately	return.	Th	e	plants	and	animals	were	formed	as
this	mud	dried.	It	was	presumed	that	man	himself	was	first	shaped	like	a	fish	and
lived	in	water.	Later,	when	he	became	capable	of	terrestrial	life,	he	cast	off	his
fish-like	capsule	like	a	butterfly	comes	out	of	its	chry	salis	and	assumed	human
form.	The	theory	is	crude	yet	the	implication	is	clear.	He	also	proposed	that
simple	forms	preceded	more	complex	forms.
3	.	Xenophanes	(576-480	BC)	contemporary	to	Anaximander,	recognised	that
fossils	are	the	remains	of	organisms	living	in	past.	According	to	him	,	the
existence	of	fossils	of	marine	animals	on	dry	land	indicated	that	the	dry	land	was
once	under	the	sea	and	that	life	originated	in	the	sea	.
4	.	Empedocles	(504-433	BC)	has	been	hailed	as	'Fathe	r	of	Evolutionary	Id	ea	'	,
by	Osborn.	He	believed	in	spontaneous	generation	and	proposed	that	evolution
of	animals	was	a	series	of	attempts	by	nature	to	produce	more	perfect	forms,	The
main	points	of	his	proposition	are	:
•	Higher	forms	of	life	evolved	gradually.
•	Imperfect	forms	(i.e.,	less	adapted	forms)	were	gradually	replaced	by	perfect
forms	(i.e.,	the	better	adapted).
•	Plant	life	came	first	and	animal	life	developed	later.
•	Perfect	forms	were	produced	by	the	extinction	of	imperfect	forms.
His	theory	was	shaped	as	follows:
All	the	matter	was	formed	of	four	elements	namely,	air,	earth,	fire	and	water.
These	were	acted	upon	by	two	great	force	s,	the	love	and	hate	,	which	caused
their	union	or	separation.	As	a	result,	parts	of	animals	were	formed	separately	as
unattached	organs.	They	joined	together	in	haphazard	manner	under	the
influence	of	love	over	hate.	The	conglomerations	produced	this	way	were	mostly
monsters	or	disharmonious	and	incapable	of	living,	but	a	few	could	function	as
successful	living	organisms.	Such	successful	combinations	populated	the	Earth.
This	theory	provides	the	first	glimmerings	of	the	idea	of	survival	of	the	fittest,
which	formed	the	basis	of	Darwin's	theory	of	natural	selection	twenty	three
centuries	later.
5.	Aristotle	(384-322	BC)	called	'the	greatest	investigator	of	antiquity',	by
Locy	(1923),	was	vitalist	and	his	ideas	dominated	biological	thoughts	well	over
a	thousand	years.	He	proposed	that	living	things	were	animated	by	a	vital	force
or	guiding	intelligence,	which	operates	constantly	and	improves	and	perfects	the
living	world.	Aristotle	suggested	that	the	various	organisms	constitute	a	series,
the	so	called	ladder	of	life	in	which	organisms	can	be	arranged	in	a	sequence	of
increasing	complexity	from	non-living	matter	through	plants	to	plant-like
animals	(like	sponges	and	sea	anemones)	or	lower	animals	and	then	to	higher
animals	.	He	placed	man	on	the	top	of	this	ladder.
Aristotle	also	introduced	the	concept	of	Teleology.	According	to	this	concept	the
natural	processes	such	as	development	or	evolution	are	guided	by	their	final
stage	or	final	goal	(Ie/os)	or	for	some	particular	purpose.	The	external	teleology
indicates	guidance	of	a	process	towards	some	specified	end	decided	by	an
external	mystical	source.	The	internal	teleology	indicates	the	end	point	of	a
process	that	has	an	understandable	materialistic	basis	that	develops	from	the
process	itself.	For	example,	plants	are	engaged	in	photosynthesis	and	animals
seek	food	for	survival	and	the	ultimate	purpose	of	survival	is	reproductive
success	.
6	.	Epicurus	(341-271	BC)	and	Soretium	(99-55	BC)	gave	an	evolutionary
explanation	of	origin	of	plants	and	animals.	Plants	appeared	before	animals	and
humans	appeared	last	of	all.
1.4.2	Pre-Darwinian	Theories
Evolutionists	of	medieval	age	were	:
I	.	Francis	Bacon	(1561-1626)	who	reviewed	Aristotelian	idea	and	presumed	that
new	species	could	arise	from	the	old	species	by	degenerative	process	caused	due
to	mutability	in	the	species.	He,	therefore,	emphasised	on	variations	as	being	the
cause	for	the	origin	of	new	species	from	the	old	one.	He	suggested	that	flying
fishes	are	intermediate	between	fishes	and	birds,	and	bats	between	birds	and
quadrupeds.	His	work	influenced	the	thinking	of	the	successors.
2	.	Jan	Swammerdam	(1637-1680),	the	Dutch	scientist,	proposed	the
'Preformation	Theory.'	According	to	this	theory	ova	contain	miniature	of	the
adult	in	preformed	state.	The	act	of	fertilisation	(i.e.,	union	with	the	sperm)
provides	initiation	for	growth	and	the	miniature	grows	into	adult.AII	parts	of	the
embryo	lie	folded	together	in	the	egg.	During	development	these	parts	grow	in
size,	and	stretch	themselves.
When	spermatozoa	were	discovered	,	they	were	called	the	animalcules	and	were
described	to	possess	the	miniature	of	the	embryo.	The	eggs	were	presumed	to
supply	nourishment	for	the	developing	embryo.
The	preformation	theory	was	discarded	by	the	valuable	observations	made	by
Casper	Friedrich	Wolf	(1759),	who	studied	chick	embryo	and	concluded	that	the
preformed	embryo	is	not	found	either	in	egg	or	sperm	.	The	development
includes	the	division	of	one	cell	and	the	modifications	in	the	cells	produced	by
its	division	to	form	various	organ	systems.
3.	Demaillet	(1656-1738)	contributed	mainl	y	on	the	nature	and	formation	of
fossils.	He	also	pointed	out	the	sim	ilarities	betw	een	aquatic	and	terrestrial	form
s	and	proposed	that	the	terres	trial	forms	have	evo	lved	from	the	marin	e	form	s
which	were	trapped	in	marsh	es.	Man	y	of	such	spec	ies	failed	to	make	the
transition	and	had	an	ill-fate.	He	cited	the	examples	of	origin	of	birds	from
flying	fish,	and	men	and	wo	me	n	from	mermen	and	merm	aid	.
4.	Maup	ertius	(	1698-	1759)	was	the	first	to	propose	a	general	theo	ry	of	evolu
tion.	He	propo	sed	that	hereditary	material	was	part	iculate	matter.	It	was	tran	sm
itted	throu	gh	both	maternal	and	patern	al	sides	of	the	famil	y.	He	thought	that
hered	itary	part	icles	co	uld	be	chan	ged	by	env	iro	nme	nt	(ac	quired	charac	ters)
.	He	a	lso	appreciated	the	role	of	natural	se	lection	in	evolution	and	of	isolation
in	spec	iation.
5.	Bonn	et	(1706-1793)	proposed	'Embo	ite	me	nt	T	heory	or	Encaseme	nt	T
heory'.	It	ad	vocated	that	the	initial	member	of	a	spec	ies	encapsulates	within	it
the	preformed	germs	of	all	future	generations.	These	ex	iste	d	insid	e	the	germ
cell	s	of	mother.	Th	e	theory	was	discredited	by	Prevost	(1824).
6	.	Wolf	Theory	of	E	p	igenesis	wa	s	proposed	by	Casper	Friedrich	Wolf	to	repl
ace	preformation	theory.	According	to	this	theory,	an	embryo	develops	by	the
gradu	al	differentiation	of	undifferentiated	simple	tissue	s	into	organs.
7	.	Linn	aeus	(	1707-	1778)	is	known	as	the	'Father	of	Taxonomy'	.	He	believed
in	specia	l	creation.	He	presumed	that	species	are	creat	ed	by	God	and	are
immutable	and	fixed	entities.
8	.	Buffon	(	1707-	1778)	be	lieved	in	the	inheritance	of	acquired	characteristics
and	the	direct	effect	of	environment	on	the	structural	modifications	of	organis
ms.	Th	ough	,	he	never	gave	a	con	sistent	theory	of	evolution	but	he	did	state
parts	of	the	theory	of	orga	nic	evolu	tion	.
9	.	Jam	es	Hutton	(1726-1797)	postulated	that	volcanic	acti	viti	es	bring	magma
up	fro	m	Earth's	molten	interior	which	on	solidification	form	s	new	igneous
rocks.	He	also	noted	that	forc	es	like	wind,	water	(rain,	surf),	heat,	cold,	ice
(glaciers)	and	acti	vities	of	plants	and	animals	erode	rocks	and	the	eroded
particles	are	transported	by	water,	and	are	deposited	in	layers.	These	layers	get
compressed	into	sedimentary	rock	s.	His	idea	of	gradual	geological	changes
brought	about	by	natural	process	is	kno	wn	as	uniformitarianism.	Thi	s	wa	s
greatly	cham	pioned	by	the	great	geologist	Charl	es	Lyell	and	has	greatly	influe
nced	Darw	in.
10	.	Era	sm	us	Darwin	(173	1-1	802),	the	grandfather	of	Charles	Robert	Darw	in,
gav	e	the	first	clear	statement	of	the	inh	eritance	of	acquired	characters,
according	to	which	the	effects	produced	by	the	environment	on	the	organi	sms
are	tran	smitted	to	the	offspring.	The	theory	wa	s	elaborated	by	Lamarck	in	the
year	1809.
T	he	contributions	made	by	Lamarck,	Darwin,	Cuvier,	Weismann,	Huxley,	etc.,
are	of	gre	at	importance,	since	the	y	pro	vok	ed	real	scientific	thinking	of
evolutionary	proc	ess	and	their	theori	es	are	still	helpfu	l,	but	in	a	somewhat
modified	forrn,	Th	e	vario	us	modem	theories	have	been	discussed	in	detail
separately,	henc	e	a	brie	f	survey	will	serve	the	purpose	here.
II	.	Lama	rc	k's	Theory	of	I	nhe	ritance	of	Acquired	Characters	(1744-1829):
Lamarck's	theory	emphasises	the	influence	of	environment	on	the	living	beings.
The	changes	introduced	by	the	environment	are	acquired	by	the	living	beings
and	are	inherited	by	the	next	generation.	Modern	supporter	of	Lamarckism	was
Lysenko	(1930),	a	Russian	botanist.
12.	Theory	of	Catastr	op	hism:	The	theory	was	formulated	to	explain	differences
in	the	past	and	present	forms	of	life	and	sharp	discontinuities	in	the	fossils
records	present	in	the	stratified	rocks.	It	state	s	that	there	had	been	several
creations,	each	preceded	by	a	catastrophe	due	to	some	supernatural	forces	and
not	the	geological	disturbances,	like	volcanic	eruptions,	upheaveling	of	Earth	,
torrential	rains	,	unprecedented	increase	in	sea	level,	etc.	Each	catastrophe
completely	destroyed	the	life.	The	new	creation	resulted	in	life	quite	different
from	the	previous	one.
George	Cuvier	(1769	-1832)	and	Orbigne	(1802-1832)	were	the	chief	advocates
of	the	Theory	of	Catastrophism.
Cuvier	(1769-1832)	is	considered	to	be	the	'Father	of	Pal	aeontology	a	nd
Comparative	Anatomy'.	Cuvier	believed	in	the	fixity	of	species.	The	occurrence
of	fossils	in	different	rock	strata	was	accounted	on	the	basis	of	catastrophism.	A
succe	ssion	of	catastrophes	have	periodically	destroyed	all	living	things,
followed	each	time	by	the	successive	creations	of	new	and	higher	forms	.
13.	Theory	of	Eternity	of	Life:	According	to	this	theory,	life	has	ever	been	in
existence	in	the	form	as	it	exists	today	and	will	continue	to	be	so	for	ever.	It
neither	had	a	beginning	nor	an	end	and	has	not	changed	or	evolved.
However,	with	present	knowledge,	the	theory	cannot	be	accepted.	The	evidences
clearly	indicate	the	gradual	complexity	in	the	organisation	of	living	beings.
14.	T	heo	ry	of	Unifor	mita	r	ia	nism:	James	Hutton	(1785)	and	Charles	Lyell
(1832)	establi	shed	the	concept	of	uniformitarianism	which	holds	that	slow	ly
acting	geological	forces	(erosion,	sedimentation,	disruption	and	uplift)	resu	lt	in
the	formation	of	fossil-bearing	rock	strata.	The	same	forces	are	acting	eve	n
today.
Jean	Baptiste	P	ierre	Antoine	de	Monet	Chevalier	de	Lamarck	(1744-1829)
known	for	'Theory	of	Inheritance	of	Acquired	Characters'
1.4.3	Evolutionary	Theories	Since	Darwin
1.	Darwin's	Theory	of	Natural	Selection	(1809-1882):	Darwin	formulated	the
theory	of	'	O	r	igin	of	Species	by	Natural	Selection	in	1859.'
To	explain	some	of	the	phenomena,	which	were	not	suitably	explained	by	natural
selection,	Darwin	propo	sed	some	more	theories.	These	are:
•	Theory	of	Pangen	esis:	To	explain	how	the	characteristics	are	transmitted	from
parent	s	to	the	offspring,	Drawin	proposed	Pangen	esis	theory.	According	to
this	theory	each	and	every	cell	of	the	body
produces	minute	primordia	called	gemmules
or	pangene.	These	gemm	ules	from	all	the
parts	of	the	body	are	carried	by	the	blood
to	the	gonads	where	these	accumulate	in	the
germ	cells.	Each	gamete	represents	minute
replica	of	parent's	body.
•	Th	eory	of	Sexual	Selection:	Darwin	pre
sumed	that	there	is	always	a	contest	among
males	for	the	possession	of	female.	For	this
reason	they	have	developed	various	methods
to	attract	the	female.	Some	are	beautifully	Charles	Robert	Darwin	(1809-1882)
co
loured,
in	othe
r
s
p
ecies	they	are	provided
who	propagated	'Theory	of	Natural	Selection'	with	horns	or	they	exhibit	different
attractive	behaviours	or	produce	sound.	This	results	in	sexua	l	dimorphism,
which	is	very	common	in	animals.
•	Ar	tificial	Selection	:	Darwin	recognised	artificial	selection	exercised	by
human	beings	as	the	commo	nest	method	for	improving	the	races	of	domestic
animals	and	cultivated	plants	and	producing	new	varieties	.	He	presumed	that	if
new	races	could	be	developed	by	artificial	selection	,	the	same	is	possible	in
nature	.
2.	'eismann's	10	I)'	of	tin	ity	f	Ger	:	Considering	the	futility	of	Darwin	's	theory	of
pangenesis,	August	Weismann	(1892),	a	staunch	supporter	of	Darwin,	proposed
that	the	cytop	lasm	of	the	anima	l	body	is	differentiated	into	soma	toplasm	and
germplasm.	The	germplasm	produces	gamates	which	transmit	the	characteristics
of	parents	into	the	offspring.	The	remain	ing	body	of	the	organ-	ism	is	formed	of
somatop	lasm.	Weismann	also	emphasised	that	only	those	changes	which	occur
in	the	germplasm	are	heritable	,	changes	occurring	in	the	body	(some	or
somatoplasm)	due	to	environmental	effect	are	not	inherited.
The	essent	ial	features	of	the	theory	can	be	summarised	as	under:
•	Ge	r	mplas	m	and	Somatoplas	m	:	Weismann	proposed	that	the	organisms	com-
prise	of	two	types	of	protoplasm-the	ger	mpla	sm	present	in	the	germ	cells	only
and	which	is	passed	on	to	the	offspring,	and	the	somatoplas	m,	the	protoplasm
forming	remainder	of	the	body	that	plays
no	role	in	heredity.	The	germ	cells	of	the
two	parents	unite	during	reproduction	and
form	the	zygote	or	fertilised	egg	.	During	development	zygote	divides	into	two
daugh
ter	ce	lls,	each	of	which	receives	an	equal	share	of	germplasm.	Through	germ
cells	a	contin	uity	of	germp	lasm	is	maintained	generation	after	generatio	n.
•	Presenc	e	of	Determinants:	Situated	in	the	germplasm	are	minute	comp	lex
structures	.
August	Weismann
(1838-1914)	proposed	'Theory	of	Continuity	of	Germplasm	'
These	are	known	as	determinants.	The	determinants	can	be	compared	with	the
present	day	chromosomes.	The	characteristics	of	the	organisms	are	represented
in	the	determinants	in	the	form	of	minute	physiological	units,	the	determiners
(equivalent	to	genes)	.
•	Immortality	of	Germplasm:	The	germplasm	is	immortal	because	it	perpetuates
from	one	generation	to	the	next	through	meiotic	division.	The	germplasm	is
maintained	generation	after	generation.	The	somatoplasm	is	mortal	and	dies	with
the	death	of	the	organism.
•	Only	those	variations	which	appear	or	which	are	introduced	in	the	germplasm
(germinal	or	heritable	variations)	can	be	inherited	and	not	those	which	appear	in
the	somatoplasm.
•	The	germplasm	is	composed	of	'ids'	i.e.,	equivalent	portions	of	germplasm
contain	all	kinds	of	determinants	present	in	the	parent	body	or	which	are
responsible	for	the	development	of	characteristics	in	the	offspring.
•	In	a	fertilised	egg	'ids'	from	both	the	parents	are	contributed	in	equal	amounts.
3.	De	Vries	T	eory	of	M	tati	n:	Darwin
in	his	Theory	of	Natural	Selection	described
the	occurrence	of	variations	but	he	did	not	ex
plain	the	method	of	their	origin.	Moreover,	he
emphasised	on	small	and	cumulative	variations.
Hugo	de	Vries	(1848-1935)	suggested	that	~....
variations	which	are	important	for	evolution	are~
sudden	and	large,	which	he	called	mutations	ort
saltations.	He	proposed	'Mutation	Theory'	in
1886	for	the	origin	of	species.	"Karl	Naegeli	and	Wanger	Gulik	emphasised	Hugo	de	Vriesthe	presence
of	some	inner	directive	which	guides	Profounded	'Mu	tation	Theory'	the	course
of	evolution	independent	of	the	environment.
4.	Reeapit	ti	T	eory	f	H	ttkel:	Ernst	Haeckel	(1811)	proposed	that	'Ontogeny
recapitulates	phylogeny',	i.e.,	the	development	of	the	individual	repeats	the
evolutionary	history	of	the	race,	condensing	some	stages	and	eliminating	the
others.
5.	ry	of	Ort	..	is:	According	to	the	theory	of	Orthogenesis	the	variations	(or	in
other	words	the	evolutionary	changes)	occur	along	certain	definite	lines,	guided
by	some	undefined	or	inherent	mystical	force.	The	term	'orthogenesis'	was
proposed	by	Haeckel	in	1893.	There	were	two	views	regarding	orthogenesis.
Karl	Von	Naegeli	believed	in	the	presence	of	some	mystical	principle	of
progressive	development	in	the	living	organisms	which	brings	about	the
particular	specialisation.	The	theory	is	merely	mythical	and	has	no	scientific
basis.	Theodor	Eimar	was	of	the	opinion	that	lines	of	evolution	are	determined
by	laws	of	organic	growth,	aided	by	inheritance	of	acquired	characters	,	and
proceed	in	specific	direction.
In	certain	cases	directional	evolution	has	resulted	to	an	enormous	increase	of
size	of	horns	which	has	ultimately	proved	to	be	harmful	to	the	organisms	and	has
led	to	their	destruction.
6.	Isolation	Theory:	The	role	of	isolation	in	evolution	was	first	emphasised	by
M.	Wanger.	He	stated	that	any	factor	or	mechanism	which	separates	the
individuals	of	a	species	into	groups,	so	that	these	are	unab	le	to	intermingle	and
interbreed,	constitutes	the	isolating	mechanism	and	is	helpful	in	the	progress	of
evolution.	It	was	supported	by	Jordan	Cellogg,	Gulick	and	Crompron.
1.4.4	Modern	Evolutionary	Theory
Modern	evolutionary	theory	has	its	foundation	in	the	Evolutionary	Synthesis	or
Modern	Synthesis	that	is	formulated	on	the	basis	of	contributions	from	Genetics,
Systematics	and	Palaeontology.	It	was	named	Neo-Darwinia	n	T	heo	ry.
The	modern	synthetic	theory	of	evolution	has	evo	lved	during	the	last	century
through	accumulation	of	facts	and	theoretical	conc	lusions	from	a	number	of
scientists	.	Theodosius	Dobzhans	ky	(1900-1975)	in	his	book	'	Ge	netics	and	th	e
O	r	igin	of	Species'	emphasised	the	role	of	genetic	changes	in	natural
populations	of	Drosophila	in	the	process	of	evo	lution.	Julian	Huxley	(1924)	and
Ernst	Mayr	(1942--43)	have	explained	the	mechanism	of	origin	of	variations	in
higher	animals	and	Stebbins	in	higher	plants.	Clevland,	Blackeslee,	Renner	and
others	have	shown	that	a	combination	of	gross	chromosomal	aberratio	ns,	rare
combinations	in	balanced	lethal	systems	and	obligate	self	fertilisation	are
important	factors	for	variation	and	evo	lution.	Rensch	(1960)	has	suggested	that
the	forces	operating	for	the	origin	of	species	also	operate	for	the	evolution	of
genera,	fami	lies	and	other	highe	r	categories.
At	present	the	synthetic	theory	of	evolution	recognises	five	basic	processes,
namely,	gene	mutations,	changes	in	chromosome	number,	genetic
recombination,	natural	selection	and	reproductive	isolation.	The	three	accessory
processes	also	contribute	to	the	evo	lutionary	phenomenon.	These	are	migration,
hybridisation	and	chance	in	small	populations.
Othniel	C.	Marsh	(	1831-	1989),	Cope	(	1840-	1897),	Mathew,	Greg	ry,	Romer
and	Simpso	n	in	America,	Woodward	(1864-1944)	and	Wa	son	in	England	and
Broom	(	1866-	1951)	in	South	Afr	ica	contrib	u	ed	imme	nsely	to	verte	brate
palaeonto	logy.	J.B.S.	Haldane	(	1892-	1964),	Fisher	(1890-1962),	and	Sewa	ll
Wright	(1889-1988)
Julian	Huxley	(1887-1975)	Ernst	Mayr	(1904-2005)	(They	explained	the
mechanism	of	orig	in	of	variation	in	higher	animals)
Ronald	A.	Fish	er	(1890-1962)	Sewall	Wright	(1889-1988)	J	.B.S.	Haldane
(1892-1964)
(They	contributed	to	the	mathematical	theory	for	change	in	gene	frequency	in
populations	under	natural	selection)
and	S.	S.	Chetverikov	(1920s)	have	provided	mathematical	theory	for	gene
frequency	change	under	natural	selection	that	leads	to	the	evo	lution	of	new
populations.	Stud-	ies	on	genus	Crepis	by	E.	B.	Babcock	provided	support	to
Neo	Darwinian	theory.
At	the	same	time,	when	evo	lutionists	were	busy	to	seek	plausible	explanation
for	evolution,	some	scientists	were	trying	to	accumulate	facts	about	evo
lutionary	process.	The	evidences	are	from	morphology,	physiology,	taxonomy
and	embryology	of	living	forms	and	the	palaeontology	(the	fossi	ls	of	previously
existing	forms)	.	The	recent	techniques	have	been	helpful	in	demonstrating	the
evolution	taking	place	in	the	laboratory	within	short	periods	of	only	a	few	years.
If	organisms	with	very	short	life	cycles,	such	as	fruit	fly	or	bacteria	are	reared
for	several	generations	in	laboratory,	new	kinds	of	indivi	duals	are	observed	in
the	progeny.	Initially,	these	indiv	idua	ls	differ	slightly	from	their	parents,	but	as
they	increase	in	number,	differences	keep	on	accumu	lating	and	a	stage	is	reac
hed	when	these	become	so	markedly	different	from	their	parents	that	they	fail	to
interbreed	with	their	parents	and	thus	form	a	new	species.
A	B
FIG.	1.3:	A.	Change	in	the	frequenc	y	of	a	gene	for	green	colour	in	beetle	that
enables	it	to	merge	with	leaf	colour;	B.	Macroevolution	.
1.4.5	Evolution	at	Different	Scales
Biological	evolution	encompasses	changes	of	vastly	different	scales:
I	.	Small-scale	Evolution:	It	includes	changes	in	the	frequency	of	genes	in	a
population	from	one	generation	to	the	next.	This	is	called	microevolution.	It
happens	on	a	small	scale	and	includes	changes	within	a	single	population.
Change	in	the	frequency	of	gene	for	dark	wings	in	beetles	from	one	generation
to	the	next	is	an	example	of	microevolution.
2	.	Large-scale	Evolution:	It	includes	the	descent	of	different	species	from	a
common	ancestor	over	many	generations.	It	is	called	macroevolution.	It	operates
above	species	level	and	leads	to	the	evolution	of	lineages.	Evolution	and
radiation	of	dinosaur	lineages,	evolution	of	horse	and	evolution	of	man	are
examples	of	macroevolution.
1.5	EVOLUTION-A	FACT	OR	JUST	A	THEORY
Concept	of	evolution	started	as	a	means	to	explain	the	phenomenon	of	a
changing	living	world	over	a	period	and	to	interpret	remarkable	similarities	in
the	living	forms	of	diverse	groups	.	It	has	now	beocme	a	well	established	fact
like	Newton's	law	of	gravitation,	because	of	supportive	observations	by
comparative	anatomists,	field	naturalists,	geologists	,	palaeontologists,
geneticists	and	biochemists.
Occurrence	of	transitional	living	forms	like	Peripatus	and	a	variety	of	fossils
support	the	view	of	gradual	evolution	of	new	forms	from	pre-existing	old
organisation.	Such	transitional	forms	are	intermediate	between	two	groups
having	characters	of	both	the	groups.	For	example:
•	Fossil	Archaeopteryx	with	its	feathers,	teeth,	claws	and	lizard-like	skeleton
shows	transition	between	reptiles	and	birds.
•	Fossil	hominids	from	Africa	with	human-like	dental	arch,	small	brain,	arms
longer	than	present	humans	but	shorter	than	modern	apes,	with	pelvis	,	feet	and
legs	for	upright	walking	support	the	view	that	man	and	apes	have	arisen	from
some	common	ancestor.
1.5.1	Major	Unsolved	Problems	of	Evolution
Despite	years	of	research	and	accumulation	of	a	lot	of	knowledge	in	various
fields	of	life	sciences,	some	challanges	to	evolutionary	science	remain
unanswered.	Some	of	these	unsolved	problems	are:
I.	Origi	f	Ufe:	How	did	living	matter	originate	from	nonliving	matter?	Was	it	a
process	that	happened	only	once	or	many	times?	Can	it	happen	today	under
natural	or	artificial	conditions?	These	are	some	of	the	questions	which	remain
unanswered.
Inspite	of	well	documented	theory	of	Biochemical	Origin	of	Life	by	Oparin	and
Haldane,	evidences	are	lacking	for	support.
2.	Origj	f	ex	:	The	following	questions	regarding	sexuality	and	sexual
reproduction	remain	unanswered:
Why	is	sexuality	so	widespread	in	nature?
•	How	did	maleness	and	femaleness	arise?
•	If	sexuality	is	so	important	for	maintaining	genetic	variability,	why	and	how
many	microorganisms	can	do	without	it?
•	How	can	we	account	for	phenomenon	like	parthenogenesis?
3.	Origin	of	Phyla:	Can	we	trace	any	relationship	between	existing	phyla	and
those	that	existed	in	the	past.	Transitional	forms	between	phy	la	are	almost
unknown	and	fossil	records	if	any	are	incomplete.	Hence,	disagreement	still
exists	about	the	number	and	relation	of	various	present	day	phyla.
4	.	Cause	of	Mass	Extinction:	Mass	extinction	has	occurred	many	times	on
Earth	since	life	originated.	However,	there	is	no	agreement	on	the	cause(s)	of
repeated	extinction	of	live	forms.	Asteroids	are	considered	to	be	one	main	reason
for	extinction,	but	are	far	from	proven	as	a	cause	for	world-wide	extinction.
Similarly,	punctuated	equilibrium	theory	is	considered	to	account	for	the	sudden
appearance	of	new	groups	and	long	persistence	of	others,	it	has	raised	many
questions	about	stability	and	extinction	of	species.
1.5.2	Importance	of	Evolution
Evolution	is	not	just	important	for	Biology,	it	is	central	to	it,	because	all	life	on
Earth	has	been	shaped	by	evolution's	key	principles	i.e.,	natural	selection	and
common	descent	with	modification.
1.	Theory	of	Evolution	provides	a	means	to	understand	:
•	the	comp	lexity	in	living	world
•	the	mechanism	of	evolution	of	resistance	to	antibiotics	in	bacteria
•	the	mechanism	of	evolutionary	adaptations
2.	Theory	of	Evo	lution	tells	us	that	we	human	beings	are	not	something
different	from	all	other	living	things,	but	we	are	the	products	of	same
evolutionary	processes	by	which	other	organisms	have	come	into	existence.
3.	Through	evo	lution	we	become	aware	that:
•	Living	forms	have	descended	fom	other	varieties	of	living	things.
•	The	organisms	that	populate	the	living	world	are	not	fixed	entities,	but	are
constantly	undergoing	modifica	tion.
KEY	TERMS
•	Catastrophism	•
•	Germplasm	Theory	•
•	Microevolution	•
•	Orthogenesis	•
•	Preformation	Theory	•
•	Theory	of	Uniformitarianism	•	Emboitement	Theory
Inheritance	of	Acquired	characters	Modern	Synthetic	Theory
Palaeontology
Somatoplasm
Theory	of	Eternity	of	Life
•	Epigenesis
•	Macroevolution
•	Mutation	Theory
•	PangenesisTheory
•	Teleology
REVIEW	QUESTIONS
I	.	Summarise	essential	feature	s	of	basic	concept	of	evolution.
2.	What	do	you	mean	by	small-scale	and	large-scale	evolutionary	changes?
3.	Define	evolution.	Who	introduced	the	term	'	evolution'?	How	does	biological
evolution	differs	from	evolution?
4.	Describe	the	contribution	of	the	follow	ing	in	the	field	of	evolution:	(a)
Aristotle	(b)	Anaximander
(c)	Ernpedocles	(d)	Aristotle
5.	Name	the	scienti	st	for	the	following	:
(a)	Father	of	Evolutionary	idea	(b)	Greatest	investigator	of	Antiquity	(c)	Father
of	Palaeontology	and	Comparative	anatomy
6.	Write	short	notes	on:
(a)	Recapitulation	theory
(c)	Theory	of	Pangenesis
7.	Explain	the	following:
(a)	Monophylectic	genealogy	(c)	Modem	evolutionary	theory	(e)	Importance	of
transitional	forms	(b)	Theory	of	Continuity	of	Germplasm	(d)	Uniformitarianism
(b)	Inheritance	of	acquired	characters	(d)	Theory	of	Orthogenesis
8.	Give	a	brief	account	of	major	unsolved	problems	of	evolution.
FURTHER	READINGS
I	.	Bowler,	Peter	J.,	1984.	Evolution	:	The	History	of	an	Idea.	Berkeley
University	of	California	Press.
2.	Dawkins,	Richard,	1986.	The	Blind	Watchmaker.	New	York:	Norton	.
3.	Ehrlich,	Paul	and	Anne,	1987.	Extinction:	The	Causes	and	Consequences	of
Disappearance	of	Species.	New	York:	Scientific	American	Books.
4.	Futuyma,	Douglas	J.,	1986.	Evolutionary	Biology	(2nd	Ed.),	Sinauer
Associates,	Inc.	Publishers	Sunderland,	Massachusetts.
5.	Gould,	Stephen	Jay,	1977.	Ever	Since	Darwin,	New	York:	Norton.
DOD
2
Evidences	for	Evolution
2.1	DOCTRINE	OF	BIOLOGICAL	EVOLUTION
E	arth	is	inhabited	by	approximately	two	million	species	of	different	kinds	of
living	beings.	They	range	from	tiny	microbes	to	giant-sized	trees,	whales	and
elephants.	Biologists	have	always	tried	to	seek	an	answer	to	the	question	of	'how
did	this	tremendous	diversity	of	life	come	to	exist	on	this	planet?'	Till	the	middle
of	nineteenth	century,	it	was	believed	that	animals	and	plants	have	arisen
spontaneously	by	specia	l	creation,	each	species	being	formed	separately.
Th	e	doctrine	of	biological	evolutio	n	which	is	now	taken	for	granted	was	based
on	the	basic	similarities	seen	in	all	living	beings,	both	in	structural	organisation
and	life	processes.	It	assumes	that	all	living	organisms	have	evolved	from	some
single	common	ancestral	form	through	the	process	of	gradual	modifications,
adaptations	and	natural	selection.	Darwin	has	described	this	phenomenon	as	'
descent	with	modification.'	The	salient	features	of	the	Doctrine	of	Biological
Evolution	are:
•	Unicellular	organisms	were	the	first	to	appear.
•	Multicellular	organisms	evolved	later	from	these	simple	unicellular	forms.
•	Early	forms	were	simple	in	structure	and	gradually	evolved	into	more	and	more
complex	forms.
•	Seed	plants	(monocot	and	dicot	plants)	in	Plant	Kingdom	and	vertebrates	in
Animal	Kingdom	are	the	last	to	evolve	from	simple	seedless	plants	and
invertebrates	respectively.
2.2	EVIDENCES	FOR	BIOLOGICAL	EVOLUTION
Doctrine	of	biological	evolution	is	supported	by	evidences	drawn	from	the	study
of	different	branches	of	biology.	These	include:
I.	Evidences	from	Comparativ	e	Anatomy	and	Morphology	(Tectology)	2.
Evidences	from	Vestigial	Organs
18	[i]	Evolutionary	Biology
3.	Evidences	from	Atavism	and	Reversion	4.	Evidences	from	Comparative
Embryology	5.	Evidences	from	Palaeontology	(Study	of	fossils)	6.	Evidences
from	Geographical	Distribution	7.	Evidences	from	Connecting	Links
8.	Evidences	from	Taxonom	y
9.	Evidence	s	from	Biochemi	stry	and	Physiology
10	.	Evidence	s	from	Molecular	Records
II	.	Evidences	from	Cyto	logy
12.	Evidences	from	Genetics
2.3	EVIDENCES	FROM	COMPARATIVE	ANATOMY	AND	MORPHOLOGY
(TECTO	LOGY)
Comparati	ve	study	of	morphology	and	anatomy	of	various	organs	and	organ
systems	reveals	that	both	similarities	as	well	as	differences	exist	in	the	structure
of	body	organs	due	to	the	similarities	and	differences	in	their	functions	.
Presence	of	basic	structural	and	functiona	l	similarities	in	the	organ	systems	of
organisms	indicates	their	common	ancestry.	It	is	illustrated	by:
I	.	Homologous	organs	(Homology)
2.	Analogo	us	organs	(Analogy	or	Homoplasy)
3.	Adaptive	diverge	nce	(Adaptive	radiation)
4.	Adaptive	convergence
O	rganisms	share	a	unity	of	plan	when	they	are	closely	related	because	of
common	descent.	This	is	substantiated	by	comparative	anatomy	and	comparative
morphology.
2.3.1	Homologous	Organs	and	Structural	Homology	(Same	Source)
Homolo	gous	organs	are	different	in	appearance	and	perform	different	functions
,	but	are	built	on	the	same	basic	pattern	and	have	a	common	origin	(same	source)
.	Therefore,	homology	is	the	similarity	in	the	basic	structure	of	organs	of
different	anima	l	groups	based	on	common	ancestry	or	origin	from	some
common	structural	patterns.	The	concept	of	homo	logy	was	introduced	by
Richard	Owen.
EXAMPLE	Homology	in	Forelimbs:	The	forelimbs	of	pterodactyl	,	bird	and	1:
bat	are	modified	for	flying,	of	dolphin,	seal	and	whale	are	modified	for
swimming,	of	sheep,	dog	and	horse	for	running	and	forelim	bs	of	man	and	shrew
for	grasping.	The	functions	of	forelimbs	in	these	anima	ls	are	entirely	different
and	so	also	their	external	"appearance.	But	these	are	constructed	on	the	same
pentadactyl	pattern	,	consisting	of	the	same	bones	(humerus,	radius-ulna,	carpals,
metacarpals	and	phalanges),	muscles	,	nerves	and	blood	vessels	arranged	on	the
same	pattern.
They	follow	th	e	same	pattern	of	development.	The	homology	can	be	explained
only	on	the	basis	that	all	of	them	have	evolved	from	the	common	ancestor.	The
differences	in	appearance	are	due	to	their	adaptations	to	carry	out	different
function	s.
Homology	can	also	be	traced	in	the	structure	of	skull,	brain	,	nerves,	muscles,
heart	and	blood	vessels	of	different	vertebrates.	(Fig.	2.1)
Dolphin
Bird	Horse
Bat	Whale	Dog	Shrew	Flying	Swimming	Running	Grasping
FIG.	2.1:	Homology	in	the	structure	of	forelimbs	in	bird	and	different	mammals.
Levels	of	Homology	in	Species
Biologists	have	recognised	homology	in	species	at	three	distinct	levels:	1.
Genetic	homology
2.	Developmental	homology
3.	Structural	homology
1.	Genetic	Homology
Genetic	homology	is	most	fundamental.	It	exists	in:
•	Similarity	in	the	DNA	sequences	found	in	different	species
•	Existence	of	universal	genetic	code,	the	same	64	codons	specify	the	same
amino	acids	from	bacteria	to	man
•	Similarity	in	the	structure	of	plasma	membrane
•	Similarity	in	the	mechanism	of	transcription	and	translation	via	same	RNA
polymerase
•	Use	of	ATP	as	energy	currency
•	Similarity	in	the	mechanism	of	DNA	replication	via	DNA	polymerase
2	.	Developmental	Homology
•	Developmenta	l	homology	is	seen	in	overall	process	of	development,	form	of
the	embryos	of	different	groups	and	fate	of	particular	embryon	ic	tissues	or
organs.	Developmental	homologies	are	due	to	homologous	genes	.
3.	Structural	Homology
•	Structural	homologies	are	similarities	in	adult	morphology	in	organisms	of
different	groups	and	are	the	results	of	homologous	genes	.
EXAMPLE	2:	Homology	in	th	e	Structu	re	of	Heart:	The	heart	is	two-chambered
in	fishes,	three-chambered	in	amphibians	and	some	reptiles,	and	four-cha
mbered	in	reptiles,	birds	and	mamma	ls.	In	fishes,	heart	has	one	auricle	and	one
ventricle.	The	auricle	receives	only	deoxygenated	blood	from	the	body	and
ventricle	sends	this	deoxygenated	blood	to	gills	for	oxygenation.	In	amphibians,
right	auric	le	receives	deoxygenated	blood	from	body	and	oxyge	nated	blood
from	lungs	and	skin.	It	gets	mixed	up	in	the	undivided	ventricle.	In	higher
reptiles	(Croco	diles),	Birds	and	Mammals,	the	oxyge	nated	and	deoxygenated
blood	are	completely	separate	d	in	left	and	right	chambers	of	heart.
We	see	that	the	structure	of	heart	in	different	groups	of	vertebrates	presents	a
gradua	l	modification	while	the	fundamental	structure	is	the	same	in	all	the
groups.	(Fig.	2.2)
Aortic	arch
J	Carotid	artery
r:...
Systemic	Ventral	Pulmonary	artery
aorta	Veins	Pulmonary	vein	Bulbous	From	body
arteriosus	Left	auricle
Ventricle
Auricle	Truncus	(Atrium)	arteriosus	Sinus
venosus
A	B
Vena	cav
Right	auricle	Pulmonary	vein
Left	auricle	Left	auricle
Left	ventricle	Left	ventricle	Right	ventricle
c	o
FIG.	2.2	:	Heart	and	blood	vessels	arising	from	it	in	the	vertebrates	series
showing	homology	in	the	ir	structure	and	distribut	ion	in:	A	.	Fish	;	B	.
Amphibian;	C.	Rep	tile	;	and	D.	Mammal.
Olfactory	lobe
Cerebellum
Medulla	oblongata
Fish	Amphibia	Reptile	Bird	Mammal
FIG.	2.3:	Homology	in	the	parts	of	brain	in	fish	,	amphib	ian,	reptile	.	bird	and
mamma	l.
EXAMPLE	3:	Homology	in	Brain	Structure:	Ranging	from	Fishes	to	Mammals,	the
brain	consists	of	similar	series	of	parts-the	olfactory	lobes,	cerebral	hemispheres,
optic	lobes	and	cerebellum	and	medulla	oblongata.	As	we	progress	through	the
series	from	Fishes	to	Mammals,	some	lobes	present	gradual	enlargement
(cerebral	hemispheres).	In	Fishes,	the	cerebral	hemispheres	are	even	smaller	than
the	optic	lobes,	but	in	Mammals	they	are	so	much	enlarged	that	they	cover	the
olfactory	lobes	in	front	and	the	optic	lobes	behind.	(Fig.	2.3)
EXAMPLE	4:	Homology	in	Insect	Mouthparts:	Mouthparts	in	insects	also	show
homology.	In	Cockroach,	these	are	modified	for	biting	and	chewing,	in
Mosquitoes	for	piercing,	in	Butterfly	for	sucking,	in	Housefly	these	are	spongy
and	adapted	for	absorbing	liquid	food.	In	all	these	cases,	mouthparts	represent
modification	of	the	basic	structure	and	have	evolved	from	the	same	prototype.
Due	to	different	feeding	habits,	some	mouthparts	are	lost	and	some	have	become
elongated	and	needle-shaped	for	piercing	the	host	skin	and	sucking	the	blood
(Mosquitoes	and	Bedbugs).
EXAMPLE	5:	Homology	in	Insect	Legs:	Legs	in	Mole	cricket,	Grasshopper,	Honey
bee,	Mantids	and	Water	beetles	are	specialised	for	digging,	jumping,	collecting
pollen,	catching	prey	and	swimming	respectively,	but	in	all	these	cases	the	legs
are	formed	of	similar	five	podomeres.
EXAMPLE	6:	Homology	in	Plant	Parts	(Leaf	Modifications):	Thoms	of
Bougainvillea	(Garden	Glory)	and	tendrils	of	Passiflora	(Passion	flower),	or
Cucurbita	are	homologous	structures.	Both	are	modified	shoots	and	are	located
in	the	axil	of	leaves.	These	axillary	branches	in	Bougainvillea	are	modified	into
thorns	for	protection	from	browsing	animals	and	in	Passiflora	and	Cucurbita	,
they	are	modified	into	tendrils	and	help	the	plant	in	climbing.	(Fig.	2.4)
Tendril
Boug	ainvillea	Passion	flower
FIG.	2.4	Homology	in	thorns	of	Bougainvillea	and
tendrils	of	Passion	flower.
S	imilarly,	ph	yllode	or	phylloclad	e	ofOpuntia	and	c1adode	ofRuscus	or
Asparagus	are	also	homologous	organs	where	stem	is	modified	for	carrying	out
photosynthesis.	These	have	different	appearance	but	are	modifications	of	stem.
(Fig.	2.5)
A	B	c
FIG.	2.5:	Homologous	structures:	A.	Phylloclade	of	Opuntia;	B.	Cladode	of
Ruscus	;	C.	Cladode	of	Aspa	ragus.
2.3.1	.1	Types	of	Homology
Homology	is	of	following	four	types:
•	Phylogenetic	hom	ology	is	similarity	among	animals	or	among	plants	of
different	species.	Homology	in	the	hand	of	man	and	forelimbs	of	horse	and	bat	is
an	example	of	phylogenetic	homology.
•	Sexua	l	homology	is	paralleli	sm	in	the	male	and	female	reproductive	organs	of
the	same	species.
•	Serial	homology	has	been	observed	among	invertebrates	.	All	arthropods	have
segmented	body	with	an	exoskeleton	of	chitin.	The	exoskeleton	is	constructed	on
the	same	basic	pattern	in	all	the	classes	of	Phylum	Arthropoda.	In	crustaceans,
all	the	segments	of	body	carry	paired	jointed	appendages.	All	of	them	are
constructed	on	a	common	structural	plan,	consisting	of	a	basal	two-segmented
portion,	the	protopodite	(coxa	and	basis)	which	bears	two	lateral	outgrowths	,	the
exopodite	and	endopodite.	The	appendages	of	various	body	segments	perform
different	functions	and	accordingly	exhibit	modifications	in	the	basic	structural
plan.	The	phenomenon	of	similarity	is	described	as	serial	homology.	(Fig.	2.6)
•	Molecular	homology	is	similarity	in	the	biomolecules	,	such	as	DNA,	the
genetic	material,	found	from	viruses	to	man.
Antennule--1
Antenna	2	o
CD
-g.	Mandible---3	!!!.
0	'
Maxillula---4	Maxilla	---5
Chelate	legs<	9
10
..::::::'11
Non-chelate	""	12
legs	13
:to	Pleopods	or	/~	:
g:	1swimmerets16
~	17S'
!!!.
18
Uropod---19
FIG.	2.6:	Appendages	of	Palaemon	showing	serial	homology,	Though	appear
drastically	different	in	appearance	they	have	same	parts-protopodite,	exopodite
and	endopodite	which	are	modified	to	carry	out	different	functions	.
Significa	nce	of	Homology	or	Homologous	Organs
Presence	of	homologous	organs	in	different	groups	confirms	:
•	common	ancestry	and	inter-relationship	among	different	groups
•	occurrence	of	divergent	evolution	,	i.e.,	the	ancestral	individuals	of	the
same	group	when	migrate	to	different	habitats,	their	organs	undergo	adaptive
modifications	in	different	environments	and	become	different
2.3.2	Analogous	Organs	and	Analogy	or	Homoplasy	('Same	form')
A	nalogous	or	homoplasious	organs	have	almost	similar	appearance	and	perform
the	same	function	but	they	develop	independently	in	totally	different	groups
through	parallel	evolution	and	are	not	inherited	from	a	common	ancestor.
Therefore,	analogy	is	the	superficial	similarity	in	appearance	between	organs	of
different	animal	groups	because	the	y	carry	out	the	same	function.	(Fig.	2.7)
2.3.2.1	Analogy	in	Animals
EXAMPLE	1:	Ana	logy	in	Win	gs:	Wings	of	an	insect	(Dragonfly),	bird	(Eagle),
mammal	(Bat)	and	reptile	(Pterodactyle)	perform	the	same	function	of	uplifting
the	body	in	the	air,	but	their	basic	structure	is	totally	different.	The	wings	of	an
insect	are	mere	expansio	ns	of	bodywall	without	any	skeletal	support	.	They	are
mere	flaps	of	chitin,	stiffened	by	a	series	of	'	veins'	.	In	Pterodactyle,	each	wing
is	an	enormous	fold	of	skin	supported	by	enlarged	fourth	finger	of	the	forelimb.
In	bird,	the	flight	surface	is	formed	by	feathers	attached	to	the	bones	of	forelimb.
In	bat,	the	wing	is	formed	of	a	fold	of	skin	and	is	called	pat	agium.	It	is
supported	with	elongated	and	outspread	phalanges	of	last	four	digits	(2nd,	3rd,
4th	and	5th	digits).	(Fig.	2.7)
EXAMPLE	2:	Analogy	in	Body	Shape:	A	fish,	an	Ichthyosaur	and	a	whale	have
stream-lined	body	and	are	adapted	for	aquatic	existence	.	But	these	belong	to
three	different	classes	of	vertebrates	with	no	traces	of	common	ancestry.	(Fig.
2.8)
Phalanges
Metacarpals	Carpals	Radius	and	ulna
A	B	FIG.	2.7:	Analogous	org	ans	:	A.	Wings	of	insect;	B.	Wing	of	bird
Metacarpals
Carpals1!:~~"'$;i.............J
Radius	Skinand	ulna	(Patagium)(fused)
Humerus	C
FIG.	2.7:	C.	Wing	of	mamma	l	(bat);	D.	Wing	of	an	extinct	reptile	(pterodactyl).
A.lchthyosaur	B.Eel
C.Fish
FIG.	2.8:	Analogy	and	adaptive	convergence	in:
A.	Ichthyosaur;	B.	Eel;	C.	Fish;	D.	Whale.
EXAMPLE	3:	Analogy	in	Fins	of	Fish	and	Flippe	rs	of	W	hale:	Fins	of	fish	and
flippers	of	whale	are	completely	unrelated	structures,	but	have	similar
appearance	and	perform	the	same	function	to	help	in	swimming.	Their	structural
details	are	totally	different.	(Fig.	2.9)
2.3.2.2	Analogous	Structures	in	Plants
EXAMPLE	1:	Potato	and	Sweet	potato	have	similar	tuberous	appearance	due	to
storage	of	food,	but	Potato	is	stem	and	Sweet	potato	is	root.
EXAMPLE	2:	Cladode	of	Ruscus	or	Asparagus	are	analogous	to	leaves	of	other
plants.	Cladode	looks	like	leaves	and	carries	out	photosynthesis	like	leaves	but	is
modified	stem.
E.	AMPI.	.	3:	Tendrils	help	in	climbing,	but	they	have	different	origins.	They	are
modified	stipules	in	Smilax,	petiole	in	Clematis,	leaflets	in	Pea,	whole	leaf	in
Wild	pea	and	axillary	buds	in	Passiflora.
Analogous	organs	are	developed	in	the	evolutionary	process	through	adaptations
of	distantly	related	organisms	to	the	same	mode	of	life.
Pterygiophores
Phalanges
Fin	rays
A.	Fin	of	a	shark	B.	Flipper	of	a	whale
FIG.	2.9:	Analogy	in	the	fin	of	A.	Shark;	B.	Flipper	of	whale.
B	c	D	EA
F	IG.	2.10:	Analogous	tendrils	developing	from	different	parts	in	plants:	A.	Stem
tendril	(from	axillary	bud)	in	Passiflora	,	B.	Leaf	tendril	(from	leaf)	in	Lathyrus;
C.	Petiole	tendril	(from	petiole	of	leaf)	in	Clematis	;	D.	Stipular	tendril	(stipule
developed	into	tendril);	E.	Leaflet	tendril	(Apical	leaflets	modified	into	tendril	in
Pisum).
A	1PI.	4.	Spines	have	different	origin,	but	carry	out	the	same	function	of
protection	against	browsers	and	dessication.	Spines	in	Opuntia	and	Berberis	are
modified	leaves	but	in	Flacourtia	the	stem	and	its	branches	are	modified	into
spines.	(Fig.	2.11)
Stem	thorns
Leaf	Leafspinesspine
	--...-ItI1"t
A	B	c
FIG.	2.11	:	Analogous	organs	in	plants:	A.	Spines	of	Opuntia	(modified	leaves);
B.	Spines
of	Berberis	(modified	leaves);	C.	Branched	thorns	of
Flacourtia	cataphracta	(modified	stem)...-it.'i-i1•	.I.
Differences	between	Homologous	and	Analogous
Organs
Homologous	Organs
1	.	They	have	the	same	basic	structural	plan.
2.	They	are	found	in	closely	related	organ	isms	which	arise	from	some	common
ancestor.
3.	They	differ	in	appearance.
4.	They	are	modified	to	carry	out	different	functions.
5.	They	have	different	inte	rnal	structure	.
6.	They	lead	to	adaptive	divergence	or	divergent	evolution.
Analogous	Organs
1	.	They	have	totally	different	structural	plan.
2.	They	are	found	in	totally	unrelated	organisms.
3	.	They	have	s	imilar	appearance.
4.	They	develop	to	carry	out	the	same	function.
5.	They	have	similar	inte	rnal	structure	.
6.	They	lead	to	convergent	evolution	or	adaptive	convergence.
2.3.3	Adaptive	Divergence	and	Adaptive	Convergence
Stud	y	of	evolution	of	different	plant	and	animal	groups	rev	ea	ls	the	phenom
enon	of	adaptive	divergence	and	adaptive	convergence	influenc	ed	by	the	env
ironme	ntal	or	habitat	differences	or	similarities	.
•	Org	ani	sm	s	of	the	same	or	close	ly	relat	ed	groups	when	occupy	different	hab
itats,	they	assum	e	different	appearanc	es	and	their	homologou	s	structures
exhibit	grea	t	divergenc	e	in	the	form	and	function	.	Thi	s	is	called	adaptive
divergence.	Adapti	ve	divergence	leads	to	adaptive	radiation.	It	is	evo	lution	in
several	spec	ialised	directions	from	a	co	mmon	genera	lised	ancestral	form	.
•	Organi	sms	of	distantly	related	or	totally	unr	elated	groups	are	found	to	develop
similar	adaptations,	while	living	in	the	same	or	sim	ilar	hab	itat.	Their	ana	logous
struc	tures,	though	apparently	similar	are	constructed	on	different	basic	plan	s.
Thi	s	is	ca	lled	adaptive	convergence	or	convergent	evolution.	Both	ada	ptive
divergence	and	adapti	ve	conve	rge	nce	provide	strong	evide	nce	in	support	of
evo	lution	and	presence	of	great	biologic	al	diversity.
2.4	EVIDENCES	FROM	VESTIGIAL	ORGANS
Th	e	vestigial	or	rudimentary	organs	are	the	useless	remnants	of	structures	or
organs	which	were	prominent	and	functional	in	ancestors.	The	se	are	often
undersized,	degenerated	and	nonfunctional.
2.4.1	Vestigial	Organs	in	Man
Man	alone	possesses	nearly	100	vestigial	structures.
I	.	Vermiform	appendix	in	man	is	the	remnant	of	caec	um	which	is	large	and
functional	in	herbivorou	s	mammals.	It	contai	ns	bacteria	that	produ	ce	enzyme
cellulase	for	the	diges	tion	of	cellulo	se.
T	he	presence	of	nonfunctional	appendix	in	man	indicates	that	ancestors	of	man
(the	early	primates)	had	a	much	coarser	diet,	feedin	g	on	resistant	vegetable
matter.	But	the	descendants	changed	food	hab	its,	the	caecum	and	appendix
being	no	longer	useful	and	gradually	reduced.
2	.	Auricular	muscles	of	external	ear	are	used	in	many	mamm	als	for	collecting
sound	waves	from	the	surroundings.	Comp	lete	set	of	muscles	for	their
movements	is	present	in	the	external	ear	of	man	but	these	muscles	are
nonfunctional.
3	.	Nictitating	membrane	or	Plica	semilunaris	is	the	third	eyelid	in	the	inner
angle	of	each	eye	in	man	and	many	mammals.	It	corresponds	to	the	nictitating
membrane	but	it	is	compl	etely	unstretchable	and	nonfunctional.
4	.	Vestigial	Tail	Vertebrae:	Early	embryo	of	man	possesses	an	external	tail	but	it
is	shed	off	much	before	the	adulthood	is	attained.	Rarely,	a	child	may	be	born
with	a	short	visible	tail.	In	adults	the	tail	is	represented	by	a	string	of	caudal
vertebrae,	which	constitute	the	coccyx	(tail	bone).	(F	ig.	2.12)
5	.	Lobe	of	the	external	ear	is	of	no	practical	benefit	to	man	,	although	serve	d
the	purpose	of	sound	gathering	in	the	ancestors	of	man.
6.	Wisdom	teeth	are	the	third	pair	of	molars.	They	are	vestigial.	These	are	last	to
erupt	or	even	fail	to	erupt.
7.	Canines	in	man	are	reduce	d	due	to	taking	soft	food	and	noncarnivorous	habit.
8.	Mammary	glands	in	males	are	rudimentary.
9.	Bod	y	hair	in	human	s	are	of	no	use	and	are	vestigial	remain	s.
2.4.2	Vestigial	Organs	in	Other	Animals
Not	only	man	but	almost	all	the	plants	and	animals	possess	vestigial	organs.	A
few	of	them	are	cited	below:
I.	Vestiges	of	Hindlimbs	and	Pel	vic:	Both	Whale	and	Python	have	vestiges	of
bones	of	hindlim	bs	and	pelvic	girdle	embedded	in	the	flesh	of	abdomen.	This
shows	that	both	of	them	have	evolved	from	ance	stors	which	had	functional
hindlimb	s.	In	Snakes	these	have	disappeared	becaus	e	of	burrowing	habit	and	in
Whales	they	are	lost	because	of	aquatic	habit.
Nipples	of	mammary	glands	in
man
B
Auricular	or	ear
muscles~iiili.i:ii
Nictitating
membrane
or	plica
semilunaris
C
Segmental	muscles	of	abdomen
Pyramidal
--l-:HI.	Ileum	muscles
Rud	imentary	tail	E	F	G	FIG.	2.12:	Some	vestigial	organs	in	human	body	:	A.
Mammary	glands	,	chest	hairs	and	coccyx	;	B.	Nictitating	membrane	or	plica
semilunaris;	C.	Auricular	or	ear	muscles;	D.	Wisdom	tooth
and	pointed	canine	;	E.	Rudimentary	tail	;	F.	Abdominal	muscles;	G.	Caecum
and	append	ix.
2.	Vestigial	Wings:	Flightless	birds	(Kiwi	of	New	Zealand	and	Ostrich	of	Africa)
possess	vestiges	of	wings	supported	by	tiny	replicas	of	usual	bones	of	a	bird	's
wing.
3	.	Splint	Bones:	In	Horse	leg,	the	splint	bones	represent	the	metacarpals	of	2nd
and	4th	digits.
4.	In	animals	living	permanently	in	deep	caves,	the	eyes	are	rudim	entary	or
vestigial.
Ribs	Flightless	or	vestigial	wings	Pelvic	Vestigial	pelvicSplint	girdle	girdle	and
hindlimb	Parts	of	girdle	bone
Femur
A	B	C	o	E
FIG.	2.13:	Vestigial	organs	in	different	animals:	A.	Vestigial	pelvic	girdle	and
bones	of	hindlimb	in	Python	;	Band	C.	Vestiges	of	pelv	ic	girdle	in	Whale	;	D.
Vestigial	wings	in	flightless	bird,	Ostrich	;	E.	Splint	bones	in	the	leg	of	Horse.
2.4.3	Vestigial	Organs	in	Plants
There	are	vestigia	l	structures	in	plants	also.	For	exa	mple:
1	.	Cutin-covered	stomata	are	present	on	the	stem	s	of	cacti	plants.
2.	In	plants	related	to	Prickly	pear,	the	leaves	are	functional,	but	in	Ruscus	and
Prickly	pear	and	in	underground	stems	,	the	leaves	are	scale-like	and	vestigial.
3.	Rudimentary	stam	ens	on	some	Aspa	ragus	plants	and	nonfunctional	pistils	on
others,	actually	represent	the	vestiges	of	ancestral	monoecious	Asparagus	plants.
4.	Cycad	sperm	that	are	passively	transported	to	the	egg	cells	have	nonfunctional
flagella	.
Significance	of	Vestigial	Organs
The	occurrence	of	vestigial	structures	in	present	day	forms	indicates	that	these
structures	were	fully	developed	and	functional	in	the	ancestors	from	which	these
present	day	forms	have	evolved.	Due	to	change	in	habit,	these	structures	were
not	needed	by	the	ancestors	and	have	gradually	reduced	to	vestiges	.	The	wide-
spread	occurrence	of	vestigial	organs	provides	evidence	for	the	occurrence	of
organic	evolution.
2.5	EVIDENCES	FROM	ATAVISM	OR	REVERSION
Ata	vism	or	reversion	is	the	reappearance	of	those	ancestral	characteristics	in	an
organi	sm	or	in	the	organisms	of	a	group	,	which	do	not	occur	normally	or	which
represent	the	reminiscent	of	normal	structures	possessed	by	the	individuals	of
othe	r	groups.
Such	abnormal	structu	res	are	known	as	atavistic	cha	r	acte	rs	.	There	are	several
examples	of	reversion	or	atavism	in	man	and	other	animals.	In	such	cases
abnormal	characters	or	structures	appear	in	the	embryo	or	in	the	adu	lt,	which
were	not	present	either	in	the	parents	or	grandparents	but	were	found	in	some
remote	ancestors.
2.5.1	Examples	of	Atavism
Atavism,	of	course	not	very	common,	is	well	illustrated	by	numerous	examples:
1.	Cervical	Fistula	in	Man:	In	fishes	there	are	five	pharyngeal	pouches	which
open	to	the	exterior.	In	man,	in	the	norma	l	course	,	only	one	pharyngeal	pouch
perforates	to	form	an	opening	from	the	pharynx	to	the	exterior	in	the	form	of
external	ear	canal	and	eustachian	tube	.	But	rarely,	the	neck	may	possess	an
additional	opening	through	which	throat	or	nasal	cavity	communicates	with	the
exterior.	This	represents	the	opening	of	an	additional	pharyngeal	pouch	to	the
exterior	and	is	known	as	cervical	fistula.
2.	Tail:	Tail	is	absent	in	man,	but	occasionally	a	human	baby	is	born	with	a	short
fleshy	tail.	It	is	devoid	of	vertebrae	and	is	removed	by	surgeon	with	no	trouble.
3.	M	amma	r	y	Glands	:	Humans	posses	s	j	ust	one	pair	of	mammary	glands	in
the	pectoral	region.	The	same	condition	is	noted	in	all	the	primates	but	in	pigs	,
these	occur	in	two	rows	one	along	either	side	of	the	chest	and	abdomen.
Sometimes,	extra-mammary	glands	or	their	nipples	appea	r	in	man	.
Human	tail
Cervical	fistula
resulting	from./.-persistent~
pharyngeal	slit
A	B
4	.	Hair	on	the	Body	and	Face	in	Irish	Dogman:	Man	is	characterised	by	scanty
hair	on	the	body	and	no	hair	on	face	but	in	the	relatives	of	man	(apes)	hair	are
present	profusely.	A	man	was	born	in	Russia	with	profuse	development	of	hair
on	the	face	and	body	(Irish	dogman).
Pointed	canine	tooth
Axillary	nippleJiJE
Thoracic	nipple
Abdominal	nipple
Rudimentary	Thick	hairs
gill	slits	on	body
C	o	F
FIG.	2.14:	Atavism	in	the	human	body:	A.	Tail	in	a	human	baby;	B.	Cervical
fistula;	C.	Rudimentary	gill	slits;	D.	Mammary	glands	in	different	regions	of	the
body;	E.	Pointed	canine;	F.	Hair	on	the	body	and	face	in	Irish	dogman.
5	.	Disappearance	of	Phalanges
of	2nd	and	4th	Digits	in	Horse:Carpals	In	modem	horse	the	third	digit	isTarsals-~~..,
most	prominent	and	its	metacarpal
bears	a	hoof,	whereas	the	2nd
and	4th	digit	s	are	absent	and
their	metacarpals	are	represented
as	splint	bone	s.	Occasionall	y
a	hor	se	possesses	one	of	the	Metacarpal-3
Splint	bones---I
t	wo	splint	bones	(2nd	and	4th)Metatarsal-3	with	phalanges	and	rarely	with
a	reduced	hoof.	This	represents
the	ancestral	character	since	the
prehistoric	horses	possessed	three......---	Hoof--E.	.toes	in	each	foot	(Fig.	2.15).
6.	Homodont	Dentition	inHindlimb	Forelimb	Piscivorous	Cetaceans:	Cetaceans	FIG.
2.15:	Splint	bones,	phalanges	and	hoof	in	the	are	mamm	als,	charactersied	by
forelimb	and	hindlimb	of	horse.	heterodont	dentition.	But	piscivorous	cetaceans
possess	simple,	conical	homodont	teeth	somewhat	similar	to	predaceous	reptiles
or	some	fishes.	Their	ancestors	are	known	to	possess	heterodont	teeth	of
mammal	s.	This	could	be	considered	to	be	a	true	reversal	of	evolution	.	(Fig.
2.16).
2.5.2	Types	of	Atavism
Atavism	can	be	categorised	into	Famil	y	atavism,	Race	atav	ism	and	Atavism
teratology.
I.	Family	Atavism:	It	includes	sudden	rea	ppearance	of	a	charact	er	or	characters
in	the	offspring	afte	r	remain	ing	lat	ent	in	the	fam	ily	for	seve	ral	generations.
Th	is	phenom	enon	is	controlled	at	gene	level	and	can	be	explained	by	simple
Mendelian	laws	of	inheritance.	For	example,	app	earance	of	red	hair	in	a	child,
whose	pa	rents	and	grand-	parent	s	all	possess	bla	ck	hair,	but	the	red	hair	were
seen	in	some	members	of	the	family	several	generations	ago.	It	mean	s	the	gene
or	genes	contro	lling	red	hair	character	could	not	gain	expression	for	several
generations	due	to	some	specific	reason	and	were	transmitted	unnoticed	or	in	the
latent	condition	but	appeared	sud	denly.
2	.	Race	Atavism:	It	includes	those	cases	of	reversal	where	one	or	more	charac
ters	of	one	race	appear	in	the	indi	viduals	of	another	race.	For	example,	the
profuse	growth	of	hair	on	the	bod	y	and	face	of	Irish	dogman	and	the	presence	of
additional	mammary	glands	or	their	rudiments	in	man	are	race	atavistic	charac
ters.
3.	Atavism	of	Teratology:	Th	is	includes	the	appea	rance	in	a	race	of	such
abnorma	l	characters	which	were	norma	l	in	other	supposedly	ances	tral	races.
The	appea	rance	of	cervical	fistula	in	man,	which	actuall	y	corresponds	to	the
gill-slit,	or	the	appearance	of	externa	l	hindlimbs	in	a	humpback	whale	or	the
homodont	dentition	in	piscivorous	cetaceans	are	examples	of	teratolo	gy.
T	hese	changes	co	uld	not	be	account	ed	by	the	changes	in	the	ge	ne	pool	s	of	the
populations	but	can	be	explained	by	the	change	in	developmental	field	s.	There
are	developmental	fields	for	each	and	every	structure	in	the	developing	embryo.
For	example,	in	mammals	there	are	distin	ct	deve	lopmental	field	s	for	incisors	,
can	ines,	molars	and	premolars.	Some	physical	factors	such	as	temperature,	pH	,
or	chemica	l	factors	may	suppress	or	change	the	development	and	differenti
ation	introducing	some	change	in	the	teeth	struc	tures	.	It	means	the	environment
can	directly	affect	the	development	of	a	tra	it.
T	he	principle	of	reversibility	was	advocated	by	L.	0	0110	in	1893	and	is	now
known	as	DoUo's	law.	It	states	that	living	organisms	do	exhibit	evolutionary
irreversibility	(the	reappearance	of	ancestral	characteristics).	The	law	has	no
exceptions	and	is	rather	a	generalisation.
A	B
F	IG.	2.16:	The	evolution	of	homodont	dentition	in	piscivorous	cetaceans	from
heterodont	ancestors	is	an	example	of	atavism:	A.	Skull	of	a	primitive
creodont	carnivore;	B.	Skull	of	Zeuglodon.	an	Eocene	whale;	C.	Skull	of	a
modern	porpoise.
2.6	EVIDENCES	FROM	COMPARATIVE	EMBRYOLOGY
Early	in	the	nineteenth	century,	Von	Baer	had	not	iced	remark	able	similarity
among	vertebrate	embryos,	whose	adults	are	markedly	different.	Darwin	and
others	concluded	that	early	developmental	stages	are	more	conservative	or
evolutionari	ly	stable	than
late	stages	or	the	adults	.	Ernst	Haeckel	(1834-1919)	was	impressed	by	the
generalised	pattern	of	development	and	the	general	resemblances	between	the
embryos	of	different	groups	of	animals	.	Haecke	l	formu	lated	the	'	Rec
apitulation	Th	eory'	or	'	Bioge	netic	Law'.	It	says	"Ontogeny	recapitulates
phylogeny".	Ontogen	y	is	the	life	history	of	an	individual	starting	from	ovum	and
phylogen	y	is	the	evolutionary	history	of	the	group.	It	includes	sequence	of	adult
ancestors	which	must	have	incurred	during	the	evolution	of	the	group	of	this
individual.	It	means	an	individual	during	its	development	repeats	the	most
important	changes	which	its	ancestors	have	undergone	during	the	long	course	of
their	evolution.
Haeckel	meant	that	early	stages	of	deve	lopment	recapitulate	the	adult	ancestra	l
forms	.	The	homologies	were	traced	at	the	following	levels:
I.	Similarities	in	early	embryonic	deve	lopment	of	animals
2.	Resemb	lance	in	the	vertebrate	embryos
3.	Resemb	lance	in	invertebrate	larvae
4.	Temporary	embryo	nic	structures
5.	Development	of	vertebrate	organs
6.	Retrogressive	metamorphosis
7.	Recapitulation	Theory	and	Biogenetic	Law
2.6.1	Similarities	in	the	Early	Development	of	Animals
The	earl	y	developmental	stage	s	of	all	the	multicellular	animals	are	similar.	All
start	their	life	from	a	fertilised	egg	called	zygote	.	It	undergoes	repeated
cleavages	and	develop	s	into	morula,	blastula	and	gas	t	r	ula.	In	gastrula,	three
germinal	layers,	i.e.,	ectoderm,	mesoderm	and	endode	r	m,	are	formed	.	These
germinal	layers	give	rise	to	the	same	types	of	parts	in	all	the	animals.	Later,
development	in	different	groups	diverges.	The	nearer	the	relationship	in	the
adults,	the	greater	is	the	similarity	in	their	development.	This	support	s	the
common	ancestry	of	all	animal	s.	(Fig	.	2.17)
Blastocoel
Blastopore	Gastrula
Zygote	Eight-cell	stage	Blastocoel
Endod
erm	+-Ectoderm
x	·
Blastula	Cross	section	(hollow	ball)	of	blastula
FIG.	2.17:	Early	embryonic	stages	(up	to	gastrula	stage)	in	the	development	of	a
multi	cellul	ar	organism.
A	B	c	D	E	F	G	H
FIG.	2.18:	Similarity	in	the	early	embryos	of	some	verteb	rates:	A.	Fish;	B.
Salamander	;	C.	Tortoise;	D.	Pigeon	;	E.	Pig;	F.	Cat;	G.	Rabbit;	and	H.	Man
2.6.2	Resemblance	among	Vertebrate	Embryos
T	he	embryos	of	Fish,	Salamander,	Tortoise,	Pigeon,	pig,	Cat,	Rabbit	and	Man
during	early	stages	of	development	resemble	each	other	so	closely	that	it	is
difficult	to	distinguish	them	from	each	other.	They	all	possess:
•	Similar	head	with	rudiments	of	eyes	and	ears
•	Pharyngea	l	clefts	or	gill	clefts,	notochord	and	embryonic	tail
•	Limbs	which	develop	as	limb	buds
•	The	notochord	which	is	replaced	by	vertebral	column	in	all	the	vertebrate
embryos
Th	e	similarity	in	embryos	of	divergent	forms	of	vertebrates	indicates	their
common	ancestry,	and	the	degree	of	similarity	in	embryos	indicates	the	degree	of
evolutionary	relationship	of	adults.
2.6.3	Resemblance	in	Invertebrate	Larv	ae
Pr	esence	of	trochophore	larva	in	annelids	and	molluscs	indicates	their	ongin
from	the	same	ancestor.	Similarity	in	the	bipinnaria	larva	of	Echinodermata	and
tornaria	larv	a	of	Hemichordata	suggests	that	both	echinoderms	and
hemichordates	have	evolved	from	the	same	common	ancestor.
2.6.4	Temporary	Embryonic	Structures
The	embryos	of	certain	animals	develop	some	temporary	nonfunctional
structures	which	disappear	before	hatching	or	birth.	For	example:
I	.	Visceral	pouches	or	gill	clefts
develop	in	the	embryos	of	all
the	land	vertebrates,	but	are	not
present	in	the	adult	.	Gill	slits	are	Visceral	useful	in	fishes	because	they
livepouches	in	water	but	are	of	no	use	for	landDorsalvertebrates,	still	they	develop	in	hollow
the	embryo.	nerve	cord	2.	The	embryos	of	all	vertebratesNotochorddevelop	notochord
which	is
replaced	by	vertebral	column	in	Postanal---"
tail
adults.
3	.	Tooth	buds	develop	in	the	embryos	of	toothless	whale	and	birds,	though
adults	do	not	have	teeth.
FIG.	2.19:	Chordate	characters	that	appear	in	all	the	chordate	embryos.
Development	of	nonfunctional	structures	in	the	life	history	of	a	bird	and	whale
suggests	that:
•	Birds	and	Whale	have	evolved	from	toothed	ancestors,	and
•	Their	embryos	repeat	for	a	short	period	the	ancestral	character,	i.e.,	the
presence	of	teeth.
2.6.5	Development	of	Vertebrate	Organs
The	de	velopment	of	various	organs	like	kidneys,	gonoducts,	gonads,	heart	,
aortic	arches,	brain,	ear,	etc.	,	in	the	embryos	of	all	the	vertebrates	follows	the
same	basic	plan	and	indicates	a	common	ancestry.	For	example,	the	highly
developed	four-chambered	heart	of	birds	and	mammals	develops	as	a	two-
chambered	tube	similar	to	heart	of	fishes.	It	becomes	three-chambered	like
amphibian	heart	and	later	on	four-chambered	heart	.	Such	a	basic	plan	of
development	of	different	organs	in	all	the	groups	of	vertebrates	supports	their
common	ancestry.
2.6.6	Retrogressive	Metamorphosis
Adults	of	certain	animals	have	degenerated	features	and	do	not	show	any
resemblance	with	other	animals	of	their	group	or	any	other	group.	But,	their
larvae	have	helped	in	establishing	their	phylogenetic	relationship.	For	example:
•	Sacculina	is	a	parasitic	crustacean	that	lives	as	ectoparasite	on	crab	's	abdomen.
The	adult	has	a	sac-like	body	but	its	larva	resembles	nauplius	larva.	During
metamorphosis,	larva	undergoes	degenerative	metamorphosis	and	loses
appendages,	gills,	mouth	,	alimentary	canal	,	sense	organ	s,	etc.	The	taxonomic
position	of	Sacculina	was	established	on	the	basis	of	its	larva.	(F	ig.	2.20)
36
Iil	Evo	lutionary	Biology
Compound	eye	Antennule
•
•
•
Antenna
Root	like	processes	of	SaccuJina
Bo	dy
of
Crab
FIG.	2.20:	Saccu	lina	.	The	paras	itic	crustacean	show	ing	dege	nerated	structure.
Herdmani	a	is	an	ascidian.	Its	chordate	nature	has	been	established	from	its
larva,	which	is	free	swimming	and	possesses	all	the	three	chordate	characters.
During	metamorphosis	larva	loses	all	the	chordate	characters	and	changes	into
the	adu	lt	ascidian.	The	adult	has	a	purse-like	body	and	no	chordate	features.
(Figs	2.21A	and	B)
Neoteny:	In	some	animals	(e.g.,	axo	lotl	larva	of	Ambystoma)	the	larva	fails	to
undergo	metamorphosis.	It	develops	gonads,	attains	sexual	maturity	and	starts
reproduc	tion.	This	is	called	neoten	y	or	paedogenesis.
Retention	of	primit	ive	or	larval	features	by	adults	provides	evidence	in	favour
of	evolu	tion.	Under	specifically	favoura	ble	circumstances	natural	selection
favours	retention	of	primiti	ve	or	larval	characters.
I	ncurrent
siphon
to	mouth
Dorsal,	hollow	nerve	cord
Excurrent	Atrium	siphon
P	harynx	/'	with
numerous
slits	Anus	Tunic
-
+-
...::----:::
-.::
Intestine	Oesophagus	Stomach
FIG.	2.21	:	A	.	An	ascidia	n	adult	without	basic	chordate	features;	B.	Larv	a	of
Herdmania	showing	chordate	features.
2.6.7	Recapitulation	Theory	and	Biogenetic	Law
The	recapitulation	theory
renamed	Biogenetic	law
was	proposed	by	Von	Baer	(1828).	It	was	revised	and	by	Ernst	Haeckel	(1868).
According	to	recapitulation
theory,	every	organism	during	its	development	repeats	or	recapitulates	in	an
abbreviated	form	the	evolutionary	history	of	its	race	.	In	Haeckel	's	words,
Ontogeny	recapitulates	phylogeny.	Ontogeny	is	the	developmental	history	of	an
individual,	while	phylogeny	is	its	ancestral	history	or	the	history	of	the	race.	An
organism	repeats	its	ancestral	history	during	its	development	which	can	be
illustrated	by	the	human	development	as	follows	:
The	fertilised	egg	may	be	compared	to	the	single-celled	ancestor	of	all	the
animals	and	the	blastula	to	a	colonial	protozoan,	which	might	have	been	the
ancestor	of	all	the	Metazoa.	Gastrula	(two-layered	cup-shaped	mass	of	cells)
represents	the	Coelenterate	ancestor	and	the	embryo	with	the	development	of
mesoderm	represents	triploblastic	stage	as	in	flatworms.
The	early	human	embryo	with	a	dorsal	hollow	nerve	cord,	a	well	developed
notochord	and	a	series	of	pharyngal	clefts	represents	the	fundamental	chordate
characters.	With	the	development	of	a	piscine	heart,	paired	arotic	arches,
primitive	pronephros	and	a	tail,	it	resembles	a	fish	embryo.	Later	on,	it
resembles	reptilian	embryo,	and	finally	develops	mammalian	characteristics.
During	the	seventh	month	of	intrauterine	development	human	embryo	resembles
a	baby	ape,	being	completely	covered	with	hair	and	having	proportionately
longer	forelimbs.	It	means	embryonic	development	(ontogeny)	in	man
recapitulates	the	history	of	the	race	(phylogeny).	This	provides	support	to
recapitulation	theory.	Other	examples	that	support	recapitulation	theory	are:
•	Larva	of	Herdmania	(i.e.,	Ascidian	tadpole)	has	chordate	characters,	but	adult
Herdmania	is	without	notochord,	nerve	cord	and	tail.
•	Development	of	Frog	includes	tadpole	larva	which	is	aquatic	and	has	fish-like
characters	like	gills	,	gill	slits,	tail	with	a	tail	fin	and	lateral	line	sense	organs.
This	suggests	that	Frog	(i.e.,	amphibians)	has	evolved	from	some	fish-like
ancestor.
•	Gymnosperms	are	not	dependent	on	water	for	fertilisation,	but	flagellated
sperm
and	water	dependency	for	fertilisation	is	found	in	primitive	gymnosperms	like
Cycas	and	Gingko.	This	shows	phylogenetic	relationship	between	Gymnosperms
and	Pteridophytes.
•	Presence	of	filamentous	protonema	during	development	of	Moss	and	Fern	(that
resembles	the	filamentous	green	algae)	suggests	algal	ancestry	of	Bryophytes
and	Pteridophytes.
•	Oak	trees	from	Southern	United	States	retain	their	leaves	throughout	the	year,
whereas	oaks	from	Northern	United	States	are	deciduous	and	shed	their	leaves
during	winter.
Von	Baer's	principle	of	embryonic	differentiation	constitutes	a	better	guide	to
embryological	evidence	for	evolution.	According	to	this	principle:
•	General	characteristics	appear	in	the	development	early	and	specialised
characters	later	on.
38	~	Evolutionary	Biology
1	.	Recapitulation	is	seen	in	invertebrate	animals	also.	For	example:	Adult
insects	have	three	pairs	of	walking	legs	but	in	embryo.	each	body	segment
carries	one	pair	of	primordia	of	legs.	This	shows	that	the	insects	have	evolved
from	an	ancestor	having	segmentally	arranged	appendages.
2	.	Biochemical	recapitulation	is	also	found	in	organisms.	For	example	:	(a)
Fishes	excrete	ammonia.
(b)	Adult	frogs	and	other	amphibians	excrete	urea	while	tadpoles	excrete
ammonia	as	in	fishes	.
(c)	Birds	excrete	uric	acid	but	embryos	excrete	ammonia	first	and	urea	later
on.
•	From	the	more	general,	the	less	general	and	finally	the	specialised	characters
appear.
•	An	animal	during	development	departs	progressively	from	the	form	of	other
animals.
•	Young	stages	of	an	animal	do	not	resemble	with	the	adults	of	different	groups
but	they	resemble	with	their	embryos.
2.7	EVIDENCES	FROM	PALAEONTOLOGY	OR	PALAEOBIOLOGY
Palaeontology	is	the	study	of	fossil	remains	of	p	lants	and	animals	that	lived	in
the	past.	Fossils	(Latin;	Jossilum	=	something	dug	out)	are	actual	remains	,
traces	or	impressions	left	by	the	organisms	that	lived	in	the	past	and	got
preserved	in	the	sedimentary	rocks.	These	include	bones,	teeth,	shells	and	other
hard	parts	of	animals	or	impressions	of	plants	pressed	into	shale	or	insects
trapped	in	tree	resin.	Over	the	last	two	centuries,	palaeonto	logists	have	studied
fossils	in	Earth's	different	strata	all	over	the	world	and	have	pieced	together	the
story	of	past	life.	The	chronological	sequence	of	fossils	in	the	rock	strata
illustrates	the	sequence	of	evolutionary	events	and	has	helped	in	building	the
broad	historical	sequence	of	biological	evolution.
Study	of	plant	fossils	is	called	palaeobotan	y	and	of	animal	fossils
palaeozoology.	Leonardo	de	Vinci	(1452-1519)	of	Italy	is	called	the	'Fathe	r	of
Palaeontology'	and	Cuvier	(1800)	the	'Founder	of	Modern	Palaeontolo	gy'.
Fossil	records	provide	the	most	direct	evidence	of	evolution,	whereas	all	other
evidences	are	indirect.
2.7.1	Formation	of	Fossils
Fossils	are	formed	in	different	ways	based	on	the	environmental	conditions	.	The
fossils	may	include	origina	l	remains	of	the	hard	parts	(bones,	teeth,	shell,	etc.)
in	the	sedimentary	rocks,	petrifaction	of	hard	and	soft	parts,	carbonised	films,
molds	(impressions	of	organisms	in	rocks),	casts	(molds	filled	with	foreign
material)	and	as	actual	remains	in	peat,	amber,	asphalt	and	ice.
The	land	anima	ls	may	also	get	fossilise	d	in	amber	(hardened	resin)	,	asphalt
(hardened	tar),	volcanic	ash,	peat	bogs	and	sand	deposits	or	in	ice.
2.7.2	Types	of	Fossils
Fossils	are	of	the	following	types:
I.	Unaltered	Remains	of	Entire	Organisms:	Under	exceptionally	favourable
conditions,	the	entire	anima	l	body	gets	preserved	in	ice,	petroleum	spring,
asphalt	,	resin,	amber	and	oil-soaked	ground	.	Woolly	mammoths	from	Siberia	in
Arctic	Tundra	remain	ed	preserved	in	ice	for	thousands	of	years.	Actually,	this
area	is	described	as	'	nature's	cold	storage	or	warehouse'.
2	.	Petrified	Fossils	(Altered	Fossils):	Petrified	fossils	are	formed	by	the
replacement	of	organic	parts	of	dead	and	decaying	organisms	molecule	by
molecu	le	by	mineral	s.	The	process	is	called	petrification.	Petrified	fossils	are
formed	in	the	sedimentary	rocks	on	the	bottom	of	lakes,	rivers	or	sea	when
animals	or	plants	or	their	parts	get	buried	in	the	sediment.	The	process	of
petrification	successfully	preserv	es	the	hard	parts.	Under	very	favourab	le
conditions,	even	the	finest	details	of	soft	tissues,	like	muscles	or	other	organs	are
also	preserved	by	the	replacement	of	their	organic	material	with	mineral	s.
•	Amber	is	hardened	resin.	A	number	of	insects	and	arthropods	are	found
preserved	in	the	amber	deposits	of	Oligocene	Epoch	from	middle	Tertiary	Period
along	Baltic	Coast.
•	Asphalt	found	in	the	tar	pits	of	Rancho	La	Bera	in	Los	Angeles	(California)	has
preserved	a	number	of	birds	and	mammals.
•	Oil	soaked	ground	in	Poland	has	remains	of	complete	Woolly	Rhinoceros.
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