Learning Objectives

➢ Understands the evolution of life on earth.

➢ Gains knowledge on theories of evolution.

➢ Interprets evidences

(anatomical, embryological and

geological) of evolution.

➢ Learns the principles of

biological evolution.

➢ Understands the importance of

gene frequencies in a population.

➢ Studies the geological time scale.

➢ Understands the importance of isolating

mechanisms in speciation.

The term evolution is used to describe heritable changes in one or more

characteristics of a population of species from one generation to the other. The present state of mankind on earth is the outcome of three kinds of evolution - chemical, organic and social or cultural evolution.

Radiometric dating of meteorites yields an estimated age for the solar system and for earth as around 4.5 – 4.6 billion years. The new born earth remained inhospitable for at least few hundred millions years. At first it was too hot. This is because the collisions of the planetesimals that coalesced to form earth released much heat to melt the entire planet. Eventually outer surface of the earth cooled and solidified to form a crust. Water vapour released from the planet’s interior cooled and condensed to form oceans. Hence origin of life can be reconstructed using indirect evidences. Consequently, biologists have turned to gather disparate bits of information and filling them together like pieces of jig saw puzzle. Many theories have been proposed to explain the origin of life. Few have been discussed in this chapter.

“Each has his own tree of ancestors, but at the top of all sits probably arboreal”

Origin of life – Evolution of life forms

Theory of special creation states that life was created by a supernatural power, respectfully referred to as “God”. According to Hinduism, Lord Brahma created the Earth. Christianity, Islam and most religions believe that God created the universe, the plants and the animals.

According to the theory of spontaneous generation or Abiogenesis, living organisms originated from non-living materials and occurred through stepwise chemical and molecular evolution over millions of years. Thomas Huxley coined the term abiogenesis.

Big bang theory explains the origin of universe as a singular huge explosion in physical terms. The primitive earth had no proper atmosphere, but consisted of ammonia, methane, hydrogen and water vapour. The temperature of the earth was extremely high. UV rays from the sun split up water molecules into hydrogen and oxygen. Gradually the temperature cooled and the water vapour condensed to form rain. Rain water filled all the depressions to form water bodies. Ammonia and methane in the atmosphere combined with oxygen to form carbon-dioxide and other gases.

Coacervates (large colloidal particles that precipitate out in aqueous medium) are the first pre-cells which gradually transformed into living cells.

According to the theory of biogenesis life arose from pre-existing life. The term biogenesis also refers to the biochemical process of production of living organisms This term was coined by Henry Bastian.

According to the theory of chemical evolution primitive organisms in the primordial environment of the earth evolved spontaneously from inorganic substances and

physical forces such, as lightning, UV radiations, volcanic activities, etc.,., Oparin (1924) suggested that the organic compounds could have undergone a series of reactions leading to more complex molecules. He proposed that the molecules formed colloidal aggregates or ‘coacervates’ in an aqueous environment. The coacervates were able to absorb and assimilate organic compounds from the environment. Haldane (1929) proposed that the primordial sea served as a vast chemical laboratory powered by solar energy. The atmosphere was oxygen free and the combination of CO2, NH3 and UV radiations gave rise to organic compounds. The sea became a ‘hot’ dilute soup containing large populations of organic monomers and polymers. They envisaged that groups of monomers and polymers acquired lipid membranes and further developed into the first living cell. Haldane coined the term prebiotic soup and this became the powerful symbol of the Oparin-Haldane view on the origin of life (1924-1929).

Oparin and Haldane independently suggested that if the primitive atmosphere was reducing and if there was appropriate supply of energy such as lightning or UV light then a wide range of organic compounds can be synthesized.

Geological time scale

The duration of the earth’s history has been divided into eras that include the Paleozoic, Mesozoic, and Cenozoic. Recent eras are further divided into periods, which are split into epochs. The geological time scale with the duration of the eras and periods with the dominant forms of life is shown in Table 6.1.

The Paleozoic era is characterized by abundance of fossils of marine invertebrates. Towards the later half, other vertebrates (marine and terrestrial) except birds and mammals

ER A YEARS IN

MILLION PERIOD EPOCH

C en

oz oi

c

1 Quaternary

Recent (Holocene)

6 Pleistocene 10

Tertiary

Pliocene 15 Miocene 20 Oligocene

100 Eocene

Paleocene

M es

oz oi

c

125 Cretaceous

150 Jurassic

180 Triassic

Pa le

oz oi

c

205 Permian

230 Carboniferous

Pennsylvanian

255 Mississippian

315 Devonian

350 Silurian

430 Ordovician

510 Cambrian

Pr ec

am br

ia n

3000

Upper

Middle

Lower

Table 6.1 Geological Time Scale

appeared. The six periods of Paleozoic era in order from oldest to the youngest are Cambrian (Age of invertebrates), Ordovician (fresh water fishes, Ostracoderms, various types of Molluscs), Silurian (origin of fishes), Devonian (Age of fishes, many types of fishes such as lung fishes, lobe finned fishes and

FAUNA FLORA

Age of Mammals Angiosperms Monocotyledons

Age of Human beings

Age of Angiosperms - Dicotyledons

Human evolution

Mammals and birds

(Golden age of Reptiles) Rise of Dinesurs

Sphenopsides, Ginkgos, Gnetales, (Dicotyledons)

Herbaceous lycopods, Ferns, Conifers, Cycads

Mammal like reptiles Arborescent lycopods

Earliest reptiles Seed ferns and Bryophytes

Earliest Amphibians and abundant Echinoderms Age of fishes Progymnosperms Earliest fishes and land invertebrates Zosterophyllum

Dominance of invertebrates

Appearance of first land plants

Fossil invertebrates Origin of algae

Multicellular organisms

Appearance of eukaryotes Planktons prokaryotes

ray finned fishes), Mississippian (earliest amphibians, Echinoderms), Pennsylvanian (earliest reptiles), Permian (mammal like reptiles).

Mesozoic era (dominance of reptiles) called the Golden age of reptiles, is divided into three periods namely, Triassic (origin of egg laying mammals),

AER	YEARS INMILLI ON	PERI OD	EPO CH	FAU NA	F L OR A
cioozenC	1	Quaternar y	R e cen t(Holo cen e)	Age o f M amm a ls	Ang ios p er msMono co t y le do ns
6	Plei sto cen e	Age o f H um an b ein gs	Age o f Ang ios p er ms -Dico t y le do ns
10	Tertiar y	Plio cen e	Hu man e volut i on
15	Mio cen e	Mamm a ls a nd b ir ds
20	Olig o cen e
100	E o cen e
Pa le o cen e
icozsoMe	125	Cretaceous	(G olden a ge o f R ep t i les)R is e o f Din es ur s	Sphen opsides,Gin kg os, G net a les,(Dico t y le do ns)
150	Jurassic	Herb ace ouslyco p o ds, F er ns,C onif er s, C yc ad s
180	Triassic
icozoleaP	205	Permian	Mamm a l li ke r ep t i les	Arb ores cen tlyc opod s
230	Carboniferous	Pennsylvanian	E arlies t r ep t i les	S e e d f er ns a ndBr yo phy te s
255
Miss iss ippi an	E arlies t A mphi bi ans a ndabun d ant E chin o der ms

| 315 |Devonian |Age o f f ishes |Prog y mn os p er ms | | 350 |Silurian |E arlies t f ishes a nd l andinver tebra tes |Z os ter ophy l lum | | 430 |Ordovician |D omin ance o finver tebra tes |App e ara nce o ff ir st l and p l ants | | 510 |Cambrian |Fos si l in ver tebra tes |Or ig in o f a lgae | | niarbmacPre |3000 |Upper |Mu lt ice l lu l ar o rga ni sm s | | Middle |App e ara nce o f eu ka r yotes | | Lower |Pl an ktons proka r yotes |

Jurassic (Dinosaurs were dominant on the earth, fossil bird – Archaeopteryx) and Cretaceous (extinction of toothed birds and dinosaurs, emergence of modern birds).

Cenozoic era (Age of mammals) is subdivided into two periods namely Tertiary and Quaternary. Tertiary period is characterized by abundant mammalian fauna. This period is subdivided into five epochs namely, Paleocene (placental mammals, Eocene (Monotremes except duck billed Platypus and Echidna, hoofed mammals and carnivores), Oligocene (higher placental mammals appeared), Miocene (origin of first man like apes) and Pliocene (origin of man from man like apes).Quaternary period witnessed decline of mammals and beginning of human social life.

The age of fossils can be determined using two methods namely, relative dating and absolute dating. Relative dating is used to determine a fossil by comparing it to similar rocks and fossils of known age. Absolute dating is used to determine the precise age of a fossil by using radiometric dating to measure the decay of isotopes.

Biological evolution

Formation of protobionts

Abiotically produced molecules can spontaneously self assemble into droplets that enclose a watery solution and maintain a chemical environment different from their surroundings. Scientists call these spheres as ‘protobionts’. Liposomes are lipids in a solution that can self assemble into a lipid bilayer. Some of the proteins inside the liposomes acquired the properties of enzymes resulting in fast multiplication of molecules.

The coacervates with nucleoprotein and nutrients had a limiting surface membrane that had the characters of a virus or free living genes. Sub sequently number of genes united to form ‘proto viruses’ somewhat similar to present day

viruses. Two major cell types that appeared during this time were significant. One form of the earliest cell contained clumps of nucleoproteins embedded in the cell substance. Such cells were similar to the Monera. They are considered as ancestral to the modern bacteria and blue green algae. The other form of earliest cells contained nucleoprotein clumps that condensed into a central mass surrounded by a thin membrane. This membrane separated nucleoproteins from the cell substances. Such cells were referred to as Protista. When the natural sources of food in the ocean declined in course of time the ancestors of Monera and Protista had to evolve different methods for food procurement. These may be summarized as parasitism, saprophytism, predator or animalism and chemosynthesis or photosynthesis. When the number of photosynthetic organisms increased there was an increase in the free O2 in the sea and atmosphere.

CH4+2O2 → CO2 +2H2O 4NH3+3O2 → 2N2+6H2O

The atmospheric oxygen combined with methane and ammonia to form CO2 and free nitrogen. The presence of the free O2 brought about the evolution of aerobic respiration which could yield large amounts of energy by oxidation of food stuffs. Thus Prokaryotes and Eukaryotes evolved.

Experimental approach to the origin of life

Urey and Miller (1953), paved way for understanding the possible synthesis of organic compounds that led to the appearance of living organisms is depicted in the Fig. 6.1. In their experiment, a mixture of gases was allowed to circulate over electric discharge from an tungsten electrode. A small flask was kept boiling and the steam emanating from it was made to mix with the mixture of gases (ammonia, methane and hydrogen) in the large chamber that was connected to the boiling water. The steam condensed to form water

which ran down the ‘U’ tube. Experiment was conducted continuously for a week and the liquid was analysed. Glycine, alanine, beta alanine and aspartic acid were identified. Thus Miller’s experiments had an insight as to the possibility of abiogenetic synthesis of large amount of variety of organic compounds in nature from a mixture of sample gases in which the only source of carbon was methane. Later in similar experiments, formation of all types of amino acids, and nitrogen bases were noticed.

Evidences for biological evolution

Paleontological evidences

Paleontology is the study of prehistoric life through fossils. Fossils are described as the true witnesses of evolution or documents

Boiling water

Tungs

Spark

Flask

Gaseo (CH4 +

Water out

Conde

Cold water i

Aqueo contai compo

Tap fo withdr sampl

Fig. 6.1 Diagrammatic representation of Urey-Mi experiment

of various geological strata of evolution. Fossilization is the process by which plant and animal remains are preserved in sedimentary rocks. They fall under three main categories.

i) Actual remains – The original hard parts such as bones, teeth or shells are preserved as such in the earth’s atmosphere. This is the most common method of fossilization. When marine animals die, their hard parts such as bones, shells, etc., are covered with sediments and are protected from further deterioration. They get preserved as such as they are preserved in vast ocean; the salinity in them prevents decay. The sediments become hardened to form definite layers or strata. For example, Woolly Mammoth that lived 22 thousand years ago were preserved in the frozen coast of Siberia as such. Several human beings and animals living

in the ancient city of Pompeii were preserved intact by volcanic ash

which gushed out from Mount Vesuvius.

ii) Petrifaction – When animals die the original portion of their body may be replaced molecule for molecule by minerals and the original substance being lost through disintegration. This method of fossilization is called petrifaction. The principle minerals involved in this type fossilization are iron pyrites, silica, calcium carbonate and bicarbonates of calcium and magnesium.

iii) Natural moulds and casts – Even after disintegration, the body of an animal might leave indelible impression on the soft mud which later becomes hardened into stones. Such impressions are called moulds. The cavities of the moulds may get filled up

ten electrode

discharge

us mixture NH3 + H2 + H2O)

nser

n

us medium ning organic unds

r awing e

ller’s

by hard minerals and get fossilized, which are called casts. Hardened faecal matter termed as coprolites occur as tiny pellets. Analysis of the coprolites enables us to understand the nature of diet the pre-historic animals thrived on.

Visit any museum nearer to your school with your teacher and identify the bones of different animals including mammals. The famous Egmore Museum is in Chennai.

Evidences from comparative anatomy

Similarities in structure between groups of organisms are accepted as indicators of relationship. For example, a comparative study of the forelimbs of different vertebrates exhibits a fundamental plan of similarity in structure. These relationships can be studied under homologous organs, analogous organs, vestigial organs, connecting links and atavistic organs.

Homologous structures**

In vertebrates, comparative anatomical studies reveal a basic plan in various structures such as fore limbs and hind limbs. Fore limbs of vertebrates exhibit anatomical similarity with each other and is made of similar bones such as humerus, radius, ulna, carpals, metacarpals and phalanges.

Carpals PhalangesUlna RadiusHumerus

Whale Cat Horse

Human

BatBird

Fig. 6.2 Forelimbs of terrestrial vertebrates to show homology

Structures which are similar in origin but perform different functions are called

homologous structures that brings about divergent evolution (Fig 6.2).

Similarly the thorn of Bougainvillea and the tendrils of Curcurbita and Pisum sativum represent homology. The thorn in former is used as a defence mechanism from grazing animals and the tendrils of latter is used as a support for climbing.

Analogous structures

Organisms having different structural patterns but similar function are termed as analogous structures. For example, the wings of birds and insects are different structurally but perform the same function of flight that brings about convergent evolution (Fig. 6.3).

Carpals Ulna

Radius

Humerus

Forewing

Hindwing

Fig. 6.3 Comparison of insect and bird wing to show their analogy

Other examples of analogous organs include the eyes of the Octopus and of

mammals and the flippers of Penguins and Dolphins. Root modification in sweet potato and stem modification in potato are considered as analogous organs. Both of these plants have a common function of storage of food.

Vestigial organs

Structures that are of no use to the possessor, and are not necessary for their existence are called vestigial organs. Vestigial organs may be considered as remnants of structures which were well developed and functional in the ancestors, but disappeared in course of evolution due to their non- utilization. Human appendix is the remnant of caecum which is functional in the digestive tract of herbivorous animals like rabbit. Cellulose digestion takes place in the caecum of these animals. Due to change in the diet containing less cellulose, caecum in human became functionless and is reduced to a vermiform appendix, which is vestigial. Other examples of vestigial organs in human beings

Fig 6.4 Embryological evidences

include coccyx, wisdom teeth, ear muscles, body hair, mammae in male, nictitating membrane of the eye, etc.,

Connecting link

The organisms which possess the characters of two different groups (transitional stage) are called connecting links. Example Peripatus (connecting link between Annelida and Arthropoda), Archeopteryx (connecting link between Reptiles and Aves).

Atavistic organs

Sudden appearance of vestigial organs in highly evolved organisms is called atavistic organs. Example, presence of tail in a human baby is an atavistic organ.

Embryological evidences

Embryology deals with the study of the development of individual from the egg to the adult stage. A detailed study of the embryonic development of different forms makes us to think that there is a close resemblance during development.

The development of heart in all vertebrates follows the same pattern of development as a

pair of tubular structures that later develop into two chambered heart in fishes, three chambered in amphibians and in most reptiles and four chambered in crocodiles, birds and mammals; indicating a common ancestry for all the vertebrates,

Hence scientists in the 19th century concluded that higher animals during their embryonic development pass through stages of lower animals (ancestors). Ernst Von Haeckel, propounded the “biogenetic law or

theory of recapitulation” which states that the life history of an individual (ontogeny) briefly repeats or recapitulates the evolutionary history of the race (phylogeny). In other words “Ontogeny recapitulates Phylogeny”. The embryonic stages of a higher animal resemble the adult stage of its ancestors. Appearance of pharyngeal gill slits, yolk sac and the appearance of tail in human embryos are some of the examples (Fig. 6.4). The biogenetic law is not universal and it is now thought that animals do not recapitulate the adult stage of any ancestors. The human embryo recapitulates the embryonic history and not the adult history of the organisms.

The comparative study of the embryo of different animals shows structural similarities among themselves. The embryos of fish, salamander, tortoise, chick and human start life as a single cell, the zygote, and undergo cleavage to produce the blastula, change to gastrula and are triploblastic. This indicates that all the above said animals have evolved from a common ancestor.

Molecular evidences

Molecular evolution is the process of change in the sequence composition of molecules such as DNA, RNA and proteins across generations. It uses principles of evolutionary biology and population genetics to explain patterns in the changes of molecules.

One of the most useful advancement in the development of molecular biology is proteins and other molecules that control life processes are conserved among species. A slight change that occurs over time in these conserved molecules (DNA, RNA and protein) are often called molecular clocks. Molecules that have been used to study evolution are cytochrome c (respiratory pathway) and rRNA (protein synthesis).

Theories of biological evolution

Lamarck’s theory

Jean Baptiste de Lamarck, was the first to postulate the theory of evolution in his famous book ‘Philosophie Zoologique’ in the year 1809. The two principles of Lamarckian theory are:

i. The theory of use and disuse - Organs that are used often will increase in size and those that are not used will degenerate. Neck in giraffe is an example of use and absence of limbs in snakes is an example for disuse theory.

ii. The theory of inheritance of acquired characters - Characters that are developed during the life time of an organism are called acquired characters and these are then inherited.

The main objection to Lamarckism

Lamarck’s “Theory of Acquired characters” was disproved by August Weismann who conducted experiments on mice for twenty generations by cutting their tails and breeding them. All mice born were with tail. Weismann proved his germplasm theory that change in the somatoplasm will not be transferred to the next generation but changes in the germplasm will be inherited.

Neo-Lamarckism

The followers of Lamarck (Neo-Lamarckists) like Cope, Osborn, Packard and Spencer tried to explain Lamarck’s theory on a more scientific basis. They considered that adaptations are universal. Organisms acquire new structures due to their adaptations to the changed environmental conditions. They argued that external conditions stimulate the somatic cells to produce certain ‘secretions’ which reach the sex cells through the blood and bring about variations in the offspring.

Darwin’s theory of Natural Selection

Charles Darwin explained the theory of evolution in his book ‘The Origin of Species by Natural Selection’. During his journey around the Earth, he made extensive observations of plants and animals. He noted a huge variety and remarkable similarities among organisms and their adaptive features to cope up to their environment. He proved that fittest organisms can survive and leave more progenies than the unfit ones through natural selection.

Darwin’s theory was based on several facts, observations and influences. They are: 1. Over production (or) prodigality of

production All living organisms increase their

population in larger number. For example, Salmon fish produces about 28 million eggs during breeding season and if all of them hatch, the seas would be filled with salmon in few generations. Elephant, the slowest breeder that can produce six young ones in its life time can produce 6 million descendants at the end of 750 years in the absence of any check. 2. Struggle for existence

Organisms struggle for food, space and mate. As these become a limiting factor, competition exists among the members of the population. Darwin denoted struggle for existence in three ways –

Intra specific struggle between the same species for food, space and mate.

Inter specific struggle with different species for food and space.

Struggle with the environment to cope with the climatic variations, flood, earthquakes, drought, etc., 3. Universal occurrence of variations

No two individuals are alike. There are variations even in identical twins. Even the children born of the same parents differ in colour, height, behavior, etc., The useful variations

found in an organism help them to overcome struggle and such variations are passed on to the next generation. 4. Origin of species by Natural Selection

According to Darwin, nature is the most powerful selective force. He compared origin of species by natural selection to a small isolated group. Darwin believed that the struggle for existence resulted in the survival of the fittest. Such organisms become better adapted to the changed environment.

Objections to Darwinism

Some objections raised against Darwinism were –

• Darwin failed to explain the mechanism of variation.

• Darwinism explains the survival of the fittest but not the arrival of the fittest.

• He focused on small fluctuating variations that are mostly non-heritable.

• He did not distinguish between somatic and germinal variations.

• He could not explain the occurrence of vestigial organs, over specialization of some organs like large tusks in extinct mammoths, oversized antlers in the extinct Irish deer, etc.,

Neo Darwinism

Neo Darwinism is the interpretation of Darwinian evolution through Natural Selection as it has been modified since it was proposed. New facts and discoveries about evolution have led to modifications of Darwinism and is supported by Wallace, Heinrich, Haeckel, Weismann and Mendel. This theory emphasizes the change in the frequency of genes in population arises due to mutation, variation, isolation and Natural selection.

Mutation theory

Hugo de Vries put forth the Mutation theory. Mutations are sudden random changes that occur in an organism that is not heritable. De Vries carried out his experiments in the Evening Primrose plant (Oenothera lamarckiana) and observed variations in them due to mutation.

According to de Vries, sudden and large variations were responsible for the origin of new species whereas Lamarck and Darwin believed in gradual accumulation of all variations as the causative factors in the origin of new species.

Hugo de Vries believed that Mutations are random and directionless, but Darwinian variations are small and directional.

Hugo de Vries believed that speciation are due to Mutation and called saltation (single step large Mutation).

Salient features of Mutation Theory

• Mutations or discontinuous variation are transmitted to other generations.

• In naturally breeding populations, mutations occur from time to time.

• There are no intermediate forms, as they are fully fledged.

• They are strictly subjected to natural selection.

Modern synthetic theory

Sewell Wright, Fisher, Mayer, Huxley,Dobzhansky, Simpson and Haeckel explained Natural Selection in the light of Post-Darwinian discoveries. According to this theory gene mutations, chromosomal mutations, genetic recombinations, natural selection and reproductive isolation are the five basic factors involved in the process of organic evolution. i. Gene mutation refers to the changes in the

structure of the gene. It is also called gene/ point mutation. It alters the phenotype of an organism and produces variations in their offspring.

ii. Chromosomal mutation refers to the changes in the structure of chromosomes due to deletion, addition, duplication, inversion or translocation. This too alters the phenotype of an organism and produces variations in their offspring.

iii. Genetic recombination is due to crossing over of genes during meiosis. This brings about genetic variations in the individuals of the same species and leads to heritable variations.

iv. Natural selection does not produce any genetic variations but once such variations occur it favours some genetic changes while rejecting others (driving force of evolution).

v. Reproductive isolation helps in preventing interbreeding between related organisms.

Evolution by anthropogenic sources

Natural Selection (Industrial melanism)

Natural selection can be explained clearly through industrial melanism. Industrial melanism is a classical case of Natural selection exhibited by the peppered moth, Biston betularia. These were available in two colours, white and black. Before industrialization peppered moth both white and black coloured were common in England. Pre-industrialization witnessed white coloured background of the wall of the buildings hence the white coloured moths escaped from their predators. Post industrialization, the tree trunks became dark due to smoke and soot let out from the industries. The black moths camouflaged on the dark bark of the trees and the white moths were easily identified by their predators. Hence the dark coloured moth population was selected and their number increased when compared to the white moths. Nature offered positive selection pressure to the

black coloured moths. The above proof shows that in a population, organisms that can adapt will survive and produce more progenies resulting in increase in population through natural selection.

Artificial selection is a byproduct of human exploitation of forests, oceans and fisheries or the use of pesticides, herbicides or drugs. For hundreds of years humans have selected various types of dogs, all of which are variants of the single species of dog. If human beings can produce new varieties in short period, then “nature” with its vast resources and long duration can easily produce new species by selection.

Adaptive Radiation

The evolutionary process which produces new species diverged from a single ancestral form becomes adapted to newly invaded habitats is called adaptive radiation. Adaptive radiations are best exemplified in closely related groups that have evolved in relatively short time. Darwin’s finches and Australian marsupials are best examples for adaptive radiation. When more than one adaptive radiation occurs in an isolated geographical area, having the same structural and functional similarity it is due to convergent evolution.

Darwin’s finches

Their common ancestor arrived on the

Galapagos about 2 million years ago. During

Tr ee

fin ch

es

Ground finches

G ra

sp ing

bi lls

Probing bills

Crushing bills

Warbler finch (Certhidea olivacea)

Woodpecker finch (Cactospiza pallida)

Cactus (Geosp

Small insectivorous tree finch

(Camarhynchus parvulus)

Large insectivorous tree finch

(Camarhynchus psittacula)

Vegetarian tree finch (Platyspiza

crassirostris)

Warbler finch

C ac

tu s

ea te

r

Insect eaters

Bud eater

Parrot-like bill

Seed eaters

Fig 6.5 Darwin’s finches

that time, Darwin’s finches have evolved into 14 recognized species differing in body size, beak shape and feeding behavior. Changes in the size and form of the beak have enabled different species to utilize different food resources such as insects, seeds, nectar from cactus flowers and blood from iguanas, all driven by Natural selection. Fig. 6.5 represents some of the finches observed by Darwin. Genetic variation in the ALX1 gene in the DNA of Darwin finches is associated with variation in the beak shape. Mild mutation in the ALX1 gene leads to phenotypic change in the shape of the beak of the Darwin finches.

Marsupials in Australia and placental mammals in North America are two subclasses of mammals they have adapted in similar way to a particular food resource, locomotory skill or climate. They were separated from the common ancestor more than 100 million year ago and each lineage continued to evolve independently. Despite temporal and geographical separation, marsupials in Australia and placental mammals in North America have produced varieties of species living in similar habitats with similar ways of life. Their overall resemblance in shape, locomotory mode, feeding and foraging are superimposed upon different modes of reproduction. This feature reflects their distinctive evolutionary relationships.

Over 200 species of marsupials live in Australia along with many fewer species of placental mammals. The marsupials have undergone adaptive radiation to occupy the diverse habitats in Australia, just as the placental mammals have radiated across North America.

ground finch iza scandens)

Sharp-beaked ground finch (Geospiza difficilis)

Small ground finch (Geospiza fuliginosa)

Medium ground finch (Geospiza fortis)

Large ground finch (Geospiza magnirostris)

Mechanism of evolution

Microevolution (evolution on a small scale) refers to the changes in allele frequencies within a population. Allele frequencies in a population may change due to four fundamental forces of evolution such as natural selection, genetic drift, mutation and gene flow.

Natural selection

It occurs when one allele (or combination of alleles of differences) makes an organism more or less fit to survive and reproduce in a given environment. If an allele reduces fitness, its frequencies tend to drop from one generation to the next.

The evolutionary path of a given gene i.e., how its allele’s change in frequency in the population across generation, may result from several evolutionary mechanisms acting at once. For example, one gene’s allele frequencies might be modified by both gene flow and genetic drift, for another gene, mutation may produce a new allele, that is favoured by natural selection (Fig. 6.6).

Selection

There are mainly three types of natural

selection

i. Stabilising Selection (centipetal selection): This type of selection operates in a stable

If ‘X’ is e birds th

survivors r

Dark rock en

light grey are spotted a

by birds mo than dark

Aa

Aa Aa

Aa aa

aa aa

aa aa

AA

Fig 6.6 Natur

environment (Fig. 6.7a). The organisms with average phenotypes survive whereas the extreme individuals from both the ends are eliminated. There is no speciation but the phenotypic stability is maintained within the population over generation. For example, measurements of sparrows that survived the storm clustered around the mean, and the sparrows that failed to survive the storm clustered around the extremes of the variation showing stabilizing selection.

ii. Directional Selection: The environment which undergoes gradual change is subjected to directional selection (Fig. 6.7b). This type of selection removes the individuals from one end towards the other end of phenotypic distribution. For example, size differences between male and female sparrows. Both male and female look alike externally but differ in body weight. Females show directional selection in relation to body weight.

iii. Disruptive Selection (centrifugal selection): When homogenous environment changes into heterogenous environment this type of selection is operational (Fig. 6.7c). The organisms of both the extreme phenotypes are selected whereas individuals with average phenotype are eliminated. This results in splitting of the population into

aten by en only eproduce

vironment

beetles nd eaten re often ones

Next generation

Aa

Aa

Aa

AA AA

AA

AA

Aa

aa

aa

al selection

sub population/species. This is a rare form of selection but leads to formation of two or more different species. It is also called adaptive radiation. E.g. Darwin’s finches- beak size in relation to seed size inhabiting Galapagos islands. Group selection and sexual selection are

other types of selection. The two major group selections are Altrusim and Kin selection.

Gene flow

Movement of genes through gametes or movement of individuals in (immigration) and out (emigration) of a population is referred to as gene flow. Organisms and gametes that enter the population may have new alleles or may bring in existing alleles but in different proportions than those already in the population. Gene flow can be a strong agent of evolution (Fig 6.8).

Migration

aa

aa

aa

aa

AA AA

AA

AA

AA

Fig 6.8 Gene flow

Peak shifts in one direction

N um

be r o

f i nd

iv id

ua ls

w ith

p he

no ty

pe

Phenotypes favoured by natural selection

Peak gets higher and narrower

(a)

Fig 6.7 Operation of natural selection on different tra

Genetic drift / Sewall Wright Effect

Genetic drift is a mechanism of evolution in which allele frequencies of a population change over generation due to chance (sampling error). Genetic drift occurs in all population sizes, but its effects are strong in a small population (Fig. 6.9). It may result in a loss of some alleles (including beneficial ones) and fixation of other alleles. Genetic drift can have major effects, when the population is reduced in size by natural disaster due to bottle neck effect or when a small group of population splits from the main population to form a new colony due to founder’s effect.

Fig 6.9 Genetic drift

(b)

Medium-sized individuals

are favoured

Two peaks form

(c)

its (a) Stabilising (b) Directional and (c) Disruptive

Phenotypesfavouredby naturale selectionwith phenotypNumber of individuals	Medium-sizedindividualsare favoured

Two peaks form

| Peak gets higherand narrower |

Peak shifts inone direction

Mutation

Although mutation is the original source of all genetic variation, mutation rate for most organisms is low. Hence new mutations on an allele frequencies from one generation to the next is usually not large.

Hardy - Weinberg Principle

In nature, populations are usually evolving such as the grass in an open meadow, wolves in a forest and bacteria in a person’s body are all natural populations. All of these populations are likely to be evolving some of their genes. Evolution does not mean that the population is moving towards perfection rather the population is changing its genetic makeup over generations. For example in a wolf population, there may be a shift in the frequency of a gene variant for black fur than grey fur. Sometimes, this type of change is due to natural selection or due to migration or due to random events.

First we will see the set of conditions required for a population not to evolve. Hardy of UK and Weinberg of Germany stated that the allele frequencies in a population are stable and are constant from generation to generation in the absence of gene flow, genetic drift, mutation, recombination and natural selection. If a population is in a state of Hardy Weinberg equilibrium, the frequencies of alleles and genotypes or sets of alleles in that population will remain same over generations. Evolution is a change in the allele frequencies in a population over time. Hence population in Hardy Weinberg is not evolving.

Suppose we have a large population of beetles, (infinitely large) and appear in two colours dark grey (black) and light grey, and their colour is determined by ‘A’ gene. ‘AA’ and ‘Aa’ beetles are dark grey and ‘aa’ beetles are light grey. In a population let’s say that ‘A’ allele has frequency (p) of 0.3 and ‘a’ allele has a frequency (q ) of 0.7. Then p+q=1.

If a population is in Hardy Weinberg equilibrium, the genotype frequency can be estimated by Hardy Weinberg equation. (p + q)2 = p2 + 2pq + q2

p2 = frequency of AA

2pq= frequency of Aa

q2= frequency of aa

p = 0.3, q = 0.7 then,

p2 = (0.3)2 = 0.09 = 9 % AA

2pq = 2(0.3) (0.7) = 0.42 = 42 % Aa

q2 = (0.7)2 0.49 = 49 % aa Hence the beetle population appears to

be in Hardy- Weinberg equilibrium. When the beetles in Hardy- Weinberg equilibrium reproduce, the allele and genotype frequency in the next generation would be: Let’s assume that the frequency of ‘A’ and ‘a’ allele in the pool of gametes that make the next generation would be the same, then there would be no variation in the progeny. The genotype frequencies of the parent appears in the next generation. (i.e. 9% AA, 42% Aa and 49% aa).

If we assume that the beetles mate randomly (selection of male gamete and female gamete in the pool of gametes), the probability of getting the offspring genotype depends on the genotype of the combining parental gametes.

Hardy Weinberg’s assumptions include

No mutation – No new alleles are generated by mutation nor the genes get duplicated or deleted. Random mating – Every organism gets a chance to mate and the mating is random with each other with no preferences for a particular genotype. No gene flow - Neither individuals nor their gametes enter (immigration) or exit (emigration) the population. Very large population size - The population should be infinite in size.

No natural selection- All alleles are fit to survive and reproduce.

If any one of these assumptions were not met, the population will not be in Hardy- Weinberg equilibrium. Only if the allele frequencies changes from one generation to the other, evolution will take place.

Origin and Evolution of Man

Mammals evolved in the early Jurassic period, about 210 million years ago (mya). Hominid evolution occurred in Asia and Africa. Hominids proved that human beings are superior to other animals and efficient in making tools and culture.

The earliest fossils of the prehistoric man like Ramapithecus and Sivapithecus lived some 14 mya and were derived from ape like Dryopithecus. Dryopithecus and Ramapithecus were hairy and walked like gorillas and chimpanzees. Ramapithecus is regarded as a possible ancester of Australopithecus and therefore of modern humans. They were vegetarians (Fig 6.10).

Australopithecus lived in East African grasslands about 5 mya and was called the

Fig 6.10 Evolu

Ramapithecus HomAustralopithecus Homo habilis

Australian ape man. He was about 1.5 meters tall with bipedal locomotion, omnivorous, semi erect, and lived in caves. Low forehead, brow ridges over the eyes, protruding face, lack of chin, low brain capacity of about 350 – 450 cc, human like dentition, lumbar curve in the vertebral column were his distinguishing features.

Homo habilis lived about 2 mya. Their brain capacity was between 650 – 800cc, and was probably vegetarian. They had bipedal locomotion and used tools made of chipped stones.

Homo erectus the first human like being was around 1.7 mya and was much closer to human in looks, skull was flatter and thicker than the modern man and had a large brain capacity of around 900 cc. Homo erectus probably ate meat Homo ergaster and Homo erectus were the first to leave Africa.

Neanderthal human was found in Neander Valley, Germany with a brain size of 1400 cc and lived between 34,000 - 1,00,000 years ago. They differ from the modern human in having semierect posture, flat cranium, sloping forehead, thin large orbits, heavy brow ridges, protruding jaws and no chin. They used animal hides to protect their bodies, knew the use of

tion of Man

Homo sapiensNeanderthal mano erectus

fire and buried their dead. They did not practice agriculture and animal domestication.

Cro-Magnon was one of the most talked forms of modern human found from the rocks of Cro-Magnon, France and is considered as the ancestor of modern Europeans. They were not only adapted to various environmental conditions, but were also known for their cave paintings, figures on floors and walls.

Homo sapiens or modern human arose in Africa some 25,000 years ago and moved to other continents and developed into distinct races. They had a brain capacity of 1300 – 1600 cc. They started cultivating crops and domesticating animals.

Isolating Mechanism

Isolation is the separation of the members of a single population into sub populations so that genetic integrity of the subpopulation can be maintained. Closely related species living in the same area do not breed together; they are prevented by isolating barriers. An isolating barrier is any evolved character of the two species that stops them from interbreeding. Several kinds of isolating barriers are distinguished. The most important distinction is Prezygotic and post zygotic isolation. Prezygotic mechanisms include those which prevent two species from coming into contact. This includes ecological, seasonal, ethological and morphological. Post zygotic mechanisms are those which act after fertilization that include hybrid sterility, hybrid inviability and hybrid breakdown.

Pre-zygotic Mechanism

i. Ecological isolation or habitat isolation –

the members of the same population may be separated from one another by a differences in their habitat. For example Rana areolata occupies burrows dug by mammals and tortoises during the day and breeds in grassy shallow ponds whereas Rana grylio breeds in deep waters. Due to the difference in their habitat the two species are able to maintain their respective species identities.

ii. Seasonal isolation – In this type of isolation, difference in the breeding seasons prevents interbreeding. E.g. Toad, Bufo americanus breeds much early in the spring; whereas Bufo fowleri breeds very late in the season. They are able to maintain their species identity because of the differences in the breeding seasons.

iii. Sexual or ethological isolation/ Behavioural isolation – Prevents mating due to the difference in their sexual behavior. The species are not separated from one another either in time or in space. The mating calls of two closely related species of frogs, Hyla versicolor (grey tree frog) and Hyla femoralis (pine wood tree frog) are different which prevents interbreeding.

iv. Morphological isolation or mechanical isolation – This type of isolation is due to the differences in their external genitalia that is seen in two different species. The size difference between the toad species Bufo quercicus and Bufo valliceps, prevents their interbreeding.

v. Physiological isolation –Though mating may occur, the gametes are prevented from fertilization due to mechanical or physiological factors. E.g. The sperms of Drosophila virilis survive only for about a day when introduced into the sperm receptacle of Drosophila americana while the sperms of Drosophila americana live for a longer time.

vi. Cytological isolation – Fertilization does not take place due to the differences in the chromosome numbers between the two species, the bull frog Rana catesbiana and gopher frog Rana areolata.

Post-zygotic Mechanism i. Hybrid inviability – In this type, the sperm

enters the egg, fertilization occurs and the embryo develops into the adult but it dies before reaching maturity. In certain fishes,

frogs, beetles, even if fertilization takes place between two species, due to genetic incompatibility they do not leave any surviving offspring.

ii. Hybrid sterility – In this type, hybrids are formed due to inter specific crosses but they are sterile due to the failure of the chromosomes to segregate normally during meiosis, example Mule (inter specific cross between a horse and a donkey).

iii. Hybrid breakdown – F1 Hybrids are viable and fertile, but F2 hybrids may be inviable or sterile.

Speciation

The process by which one species evolves into one or more different species is called speciation. A.E. Emerson defines species as a ‘genetically distinctive, reproductively isolated natural population’. Speciation is a fundamental process in evolution. Evolution of a new species in a single lineage is called an agenesis / phyletic speciation. If one species diverges to become two or more species it is cladogenesis or divergent evolution.

Sympatric speciation/ Reproductive isolation

It is a mode of speciation through which new species form from a single ancestral species while both species continue to inhabit the same geographical region. Two or more species are involved. New species formed due to genetic modification in the ancestor that is naturally selected can no longer breed with the parent population. Sexual isolation is strongest. Phenotypic plasticity has emerged as potentially important first step in speciation initiated within an isolated population.

A well studied example is the adaptation of Apple maggots that feed on apples in North America. When the apple trees were imported to North America, Apple maggot flies (Rhagoletis pomonella) a parasitic insect that normally laid its eggs in the fruit of wild hawthorns until one subset of population began to lay its eggs in the fruit of domesticated apple trees (Malus

domestica) that grew in the same area. This small group of apple maggot flies selected a different host species from the rest of the population and its offsprings became accustomed to domesticated apples.

Phenotypic plasicity is the ability of single genotype to produce more than one phenotype. When this plasticity is expressed seasonally in planktons, it is referred to as cyclomorphosis.

Allopatric speciation/ Geographical speciation

It is a mode of speciation that occurs when biological populations of similar species become isolated from each other that prevents gene flow. One species becomes two species due to geographical barriers hence new species is evolved e.g. Darwin’s finches. The barriers are land separation, migration or mountain formation. When barriers occur between species, change in ecological conditions and environment leads to adaptations that produce differences. If there are no adaptations, they will not survive. Sexual isolation is weakest.

Extinction of Animals

Extinction

Extinction was common if not inevitable because species could not always adapt to large or rapid environmental changes. The impact of extinction can conveniently be considered at three levels.

Species extinction eliminates an entire species, by an environmental event (flood etc.,) or by biological event (disease or non availability of food resource half or more).

Mass extinction eliminates half or more species in a region or ecosystem, as might occur following a volcanic eruption. Five major mass extinction that occurred since the Cambrian period. This mass extinction is often referred to as K-T extinction. Table 6.2 represents the K-T extinction.

Table 6.2 Details of the five major mass extinctions since the Cambrian period

Extinction Period Date (Mya)

Estimated % of Marine Animal Extinction Genera Species

End of the Ordovician

440 61 85

End of the Devonian 365 55 82 End of the Permian 245 84 96 End of the Triassic 208 50 76 End of the Cretaceous 65 50 76

K-T Extinction refers to the German word Cretaceous and Tertiary periods.

Global extinction eliminates most of the species on a large scale or larger taxonomic groups in the continent or the Earth. Snow ball Earth and extinction following elevation in CO2 levels are example. Extinction events opens up new habitats and so can facilitate the radiation of organisms that survived the mass extinction.

Summary Evolutionary Biology is the study of history

of life forms on Earth which originated on Earth millions of years ago. How Earth originated, how life originated, what is the place of man in the universe are all general questions. This chapter deals with several theories to explain the life on Earth. Evidence from the fossil record and many other areas of biology like embryology, anatomy and molecular biology indicates a common ancestry.

The theories advanced by Lamarck, Darwin, Hugo de Vries explained the intricate evolutionary process. Geological time scale with different eras, periods and epochs gives an idea about the dominant species in those days. The mathematical distribution of gene and genotype frequencies remains constant in a small population was contributed by Hardy and Weinberg in 1608. Natural Selection and gene pool are the important factors those affect Hardy Weinberg equilibrium.

Human evolution states that humans developed from primates or ape like ancestors. The emergence of Homo sapiens as a distinct species from apes and placental mammals in brain size, eating habit and other behavior proves that ‘Ontogeny recapitulates Phylogeny’.

Separation of a single population into its subunits preserves the genetic integrity. Types of isolating mechanism and speciation prevent interbreeding of different species. Various causes of extinction of animals and the levels of impact are dealt with in the lesson.

Evaluation 1) The first life on earth

originated a) in air b) on land c) in water d) on mountain

2. Who published the book “Origin of species by Natural Selection” in 1859? a) Charles Darwin b) Lamarck c) Weismann d) Hugo de Vries

3. Which of the following was the contribution of Hugo de Vries? a) Theory of mutation b) Theory of natural Selection c) Theory of inheritance of acquired

characters d) Germplasm theory

4. The wings of birds and butterflies is an example of a) Adaptive radiation b) convergent evolution c) divergent evolution d) variation

5. The phenomenon of “ Industrial Melanism” demonstrates a) Natural selection b) induced mutation c) reproductive isolation d) geographical isolation

6. Darwin’s finches are an excellent example of a) connecting links b) seasonal migration c) adaptive radiation d) parasitism

Extinction Period	Date(Mya)	Estimated% of MarineAnimalExtinction
Genera	Species
End of the Ordovician	440	61	85
End of the Devonian	365	55	82
End of the Permian	245	84	96
End of the Triassic	208	50	76
End of the Cretaceous	65	50	76

7. Who proposed the Germplasm theory? a) Darwin b) August Weismann c) Lamarck d) Alfred Wallace

8. The age of fossils can be determined by a) electron microscope b) weighing the fossils c) carbon dating d) analysis of bones

9. Fossils are generally found in a) igneous rocks b) metamorphic rocks c) volcanic rocks d) sedimentary rocks

10. Evolutionary history of an organism is called a) ancestry b) ontogeny c) phylogeny d) paleontology

11. The golden age of reptiles was a) Mesozoic era b) Cenozoic era c) Paleozoic era d) Proterozoic era

12. Which period was called “Age of fishes”? a) Permian b) Triassic c) Devonian d) Ordovician

13. Modern man belongs to which period? a) Quaternary b) Cretaceous c) Silurian d) Cambrian

14. The Neanderthal man had the brain capacity of a ) 650 – 800cc b) 1200cc c) 900cc d) 1400cc

15. According to Darwin, the organic evolution is due to

a) Intraspecific competition b) Interspecific competition c) Competition within closely related species. d) Reduced feeding efficiency in one species due to the presence of interfering species.

16. A population will not exist in Hardy- Weinberg equilibrium if

a) Individuals mate selectively b) There are no mutations c) There is no migration d) The population is large

17. List out the major gases seems to be found in the primitive earth.

18. Explain the three major categories in which fossilization occur?

19. Differentiate between divergent evolution and convergent evolution with one example for each.

20. How does Hardy-Weinberg’s expression (p2+2pq+q2=1) explain that genetic equilibrium is maintained in a population? List any four factors that can disturb the genetic equilibrium.

21. Explain how mutations, natural selection and genetic drift affect Hardy Weinberg equilibrium.

22. How did Darwin explain fitness of organisms?

23. Mention the main objections to Darwinism. 24) Taking the example of Peppered moth,

explain the action of natural selection. What do you call the above phenomenon?

25. Darwin’s finches and Australian marsupials are suitable examples of adaptive radiation – Justify the statement.

26. Who disproved Lamarck’s Theory of acquired characters? How?

27. How does Mutation theory of De Vries differ from Lamarck and Darwin’s view in the origin of new species.

28. Explain stabilizing, directional and disruptive selection with examples.

29. Rearrange the descent in human evolution Austrolopithecus → Homo erectus → Homo sapiens → Ramapithecus → Homo habilis.

30. How does Neanderthal man differ from the modern man in appearance?

31. Define isolating mechanism and explain its types with suitable examples.

32. Define speciation according to A.E. Emerson and explain its types giving suitable examples.

33. Give an account on the major causes for the extinction of a particular species on earth.

34. Explain the three level of impact of extinction of species.

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ce pt

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