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Approaches to the Study of Behaviour 29 these and similar social systems in birds reveal that nests with helpers are more successful, i.c., the number of young fledglings is higher. In fact, the various theories and concepts that constitute sociobiology have their roots in many earlier works. Among the most significant are the writings of Williams (1966) on natural selection and concept of adaptation, Trivers (1971, 1972) on the evolutionary aspects of altruism and parental behaviour and Hamilton (1964, 1971) on the genetic theory underlying the evolution of social behaviour, Studies conducted under the general heading of sociobiology include, for example, those on altruism in ground squirrels (Sherman, 1977); on strategies for reproduction in damselflies for reproduction in damselflies and other insects (Waage, 1979, 1997); on parental investment in water bugs (Smith, 1997) and on mate slection in American kestrels or European falcons (Duncan and Bird, 1989). In recent years, a major topic for investigators using the sociobiological approach has involved sexual selection and various factors, influencing mate choice (Anderson, 1994; Cowaty, 1995; Eberhard, 1996). M. Petrie et al., (1991) reported that peahens prefer peacocks with elaborate trains (i.e., greater number of eyespots in their tail feathers). Prasad and Chaturvedi (2006) studied 12-hour temporal relationship of circadian serotonergic and dopaminergic activity influences seasonal testicular growth and secondary sex characters in Indian weaver bird, Ploceus phillippinus. 2.2, TOOLS AND TECHNIQUES OF ANIMAL BEHAVIOUR The discipline of ethology may be approached from different viewpoints depending upon the questions asked and the inclination of the investigator. This is in part a reflection of the diversity of the animals, the tools available for study, the diversity in behaviour and an accident of history. The following three disciplines have converged to produce its current complexion: 1. Psychological techniques. Psychologists have been traditionally interested in questions involved with how an animal learns — its motivation and drive to perform certain acts. Thus they devise puzzle-boxes, skinner-boxes and mazes and training tasks for animals as their methodological approach. 2. Physiological techniques. Physiologists have been traditionally interested in describing the sense organ response, the function of nervous system and effect of hormones and pheromones on the animal behaviour. They record the sens electronic equipments. For example, neurophysiology of cricket song was studied with the aid of implanted micro-electrodes. Likewise, to understand the magnetic sense of pigeons used during homing and their migration can be studied by the help of Helmholtz coils which may be set above the pigeon’s head and around its neck. ‘The psychological techniques and physiological techniques are performed in the laboratory (see Section 2.3). 3. Field biological techniques. Field biologists have traditionally been inter animal's relationship with its environment — the social behaviour and behavioura that permit individual and population survival. They often use observatior techniques perhaps aided by camera, radar, radio transmitters, oscilloscop flash lamp, ultrasonic loudspeakers, multipen event recorder and healthy am Field biologists compare behavioural patterns between individuals and bet may expose the animal to different environments to know how behaviour done to understand the relevance of the behaviour to the animal’s past and Certain significant field-biological tools and techniques of ethology ar 1. Study of animal’s behaviour in its natural surrounding. The firs observe the behaviour of an animal in general in as many contexts as possibh that the surroundings can distort and modify behaviour, the study should be mad which are as natural as possible, Ideally we would follow the animal around of in the wild state. In practice we usually have to compromise. A common sol 30 ‘Animal Behaviour animal as comfortable as possible and allow il to behave in as natural fashion as it will when lef without disturbance, The bulk of behaviour data is gathered in this way, and it is supplemented by whatever field observations are possible, Gradually a picture is constructed of the motor patterns which the animal uses in its daily life, the stimuli (both physical and social) to which it seems to be responsive and the ways in which the behaviour changes witha shift in the physiological condition of the animal. A camouflage tent is also an aid in observing the natural behaviour of animals which soon get used to it. In this way the scientist can make his observations without being noticed by the subject (Fig. 2.3). 2, Selection of experimental animal. The success ofa behavioural investigation depends very much on the selection of an experimental animal (subject) which is appropriate to the area of interest. Obviously a study of birds is likely to be a poor introduction to the use of olfaction in social communication. 3. Quantification of behaviour. To gain full picture of an animal's behaviour it is often necessary to record its activities, which are frequently repetitive, over a long period of time, To do this by direct observation would drive all investigators insane; therefore a variety of activity recorders are used. For example, multi-pen recorders allow continuous recording of an animal's behaviour. Each pen of the recorder is wired to a separate switch of the manual keyboard. When a behaviour occurs, the appropriate key is depressed to actuate (to move to action) the recording pen and the key is held down until the behaviour terminates. The duration of each behaviour is also recorded by stop-watch or other means, The recordings provide a permanent record of the observations. ‘A time-sampling method may improve the accuracy of behavioural observation. This method uses a timing device to pace the observer's recording. For example, a tape-recorded signal 23. A traditional view of the distinction between delivered through an earphone to the q " 7 cthol y. The psycholc its hi observer marks off fifteen-serond oe eee age unt i's Nh snp yoo sm pthe etholog selFin the box observer notes the behaviours f he i being displayed. intervals and on hearing each signal fr slow-moving animals such a ‘Salticid spiders, multiple- ‘Approaches to the Study of Behaviour 3a 5. Use of video-tape in analysis of behaviour. The current use of audio-visual media has revolutionized the science of animal behaviour, Video-taping fills the need for a visual record of behaviour as it actually occurred. By repeated analysis of the same behavioural sequence the investigator often makes critical observations.not seen at the first time, In addition, repeated data collection on the same behavioural sequence provides the observer reliability of the data estimation. Video-taping has several advantages over filming: 1. The visual record is available for use immediately after taping, because there is no processing delay, 2. The video monitor enables one to view the visual record while taping is actually in progress, thus enabling mechanical errors to be rectified immediately, 3. Taping may be done under normal lighting conditions, thereby avoiding bright lights and intense heat from interfering with the subjects’s behaviour, 4, Because video-tape may be reused, it is less expensive means of recording behaviour once the initial equipment has been purchased, 5. Video-tapes are more suitable than film for recording Jong sequences of behaviour since they are capable of longer continuous recording, 6. Video- tape provides a more convenient means of linking vocalisation (auditory signals with behavioural patterns), ‘Aside from the initial cost, there are two major disadvantages to video-tape equipment. It is generally less portable than movie cameras, thus, confining taping techniques largely to laboratory studies. Unlike movie film, video-tape is not easily sectioned into single-frame visual records to be reproduced as slides and photos, or made into line drawings. 6. Use of tranquilizers and radio-telemetry. To study social behaviour, observations of natural behaviour in individuals (wild animals) over long periods of time become important. The tranquilizer gun enables us to capture animals, mark them and release them unharmed. For this a shell is filled with an anesthetic drug, then loaded into an air rifle and shot at the animal and then the shell with its barbed hook is removed. The scientists now have upto an hour to take measurements and to mark the subject. One method of marking which allows us to track the individual animal more easily is radio-telemetry. In this method, an anesthized animal (e.g., lions) is fitted with a collar containing batteries and a radio transmitter. A receiver to make the transmitter signals audible ‘and a directional antenna enable us to locate the animal at any time, even in dense under bush. 2.3. STUDY OF BEHAVIOUR IN LABORATORY ‘The first major goal of brain researchers or psychologists was the localization of brain functions, to know which areas or structures of the brain were involved in controlling particular behaviour. These studies include following three types of techniques. ‘A. Neuroanatomical or Lesion or Ablation technique B, Neurophysiological technique C. Neurochemical technique ‘A.Neuroanatomical or Lesion or Ablation Techniqui This is oldest and most crude method for studying neural structure and behaviour. For this purpose, certain. their functions is deduced from the abnormal behavioul ablation (i.e., removal of tissue) may be carried out fre the localised and small lesions (i.e., pathological char electric currents through platinum, irridium electrode to also be created by neurotoxic kainic acid. Lesion method in the latter case, neurons are activated while in the Neuroanatomical technique helped many workers to fo are known stereotaxic atlases that are presently availa rat (De Groot, 1959), cat (Jasper and Marsan, 1954)¢ yo 5 32 Animal Bohaviou streutures such as amygdala influence aggressive ang ed by lesion experiments, Bilateral lesions in amygday ations that brai defence behaviour in cats was obtai caused enhanced sexual behaviour. Pierre-Paul Broca (1824-1880) was a French doctor who in 1861 came across a patie; with head injury whose speech was defective, This correlation of brain lesion and fae viscinated him and he started observing different patients with similar injuries. After few years, roca concluded that there is a definite area in the brain cortex which is responsible the articulate speech, Later using the same technique another scientist, a German neurologist, Carl Wernicke (1848-1905), in 1880, established another arca in the brain responsible for speech. These two «reas are known as Broca’s and Wernicke’s areas (Fig. 2.4), respectively. Broca’s area controls ig. 2.4. Human brain showing Broca’s and Wernicke’s areas for speech. the infrastructure for speaking a word, this means it has control over neck muscles and larynx. Wernicke’s area is word retriever or dictionary. A lesion in Broca’s area causes slurring in speech and there is difficulty in understanding the words spoken by such a pero” though the person would talk sensibly. Opposite to it, if there is a lesion in Wernicke’s area then the person tan talk clearly but the speech will not be articulated or coherent. ‘The toads (amphibians) have, been the, major objects for ablation techniques: the experimental finding that there are zones in different areas of the toad brain responsible for the triggering of prey catching (Optie teetuim) and avoidance behaviour egion), is supported tie iat cages 2 i by ablation experiments, Different nuclei of hyp g olling vital activities such as feeding, drinking, discovered by lesion technique. Lesion in hy yperphagia (ie., abnormally increased th (ie., abnormally increased aj entromedial hypothalamus increases hua this area of hypothalamus has Uices hunger and animal stop eating. It -_— ‘Approaches to the Study of Behaviour 33 By further findings, it has also been established that there are specific nuclei in the anterior and ventro-medial parts of hypothalamus that start and stop the sexual behaviour. Dorsal area Paraventricular. Lateral nucleus hypothalamus Posterior hypothalamus, - Lateral preoptic nucleus Dorsomedial nucleus Mamilary Medial preoptic body nucleus ‘Supra optic nucleus Ventromedial Optic nerve nucleus: Anterior’ hypothalamus Jnfundibulum Pituitary Fig. 2.5. Nuclei of hypothalamus of human brain. 2, Neurophysiologieal Technique The discovery of animal electricity or bioelectricity came by chance. The Italian scientist, Luigi Galvani (1737-1798), in 1786, hung some frog legs on his balcony railing one day and noticed that the frog’s legs twitched when they touched the metal fence. Galvani at first misinterpreted this phenomenon attributing it to muscle alone. By 1793, Galvani discovered that twitches occurred in the nerve muscle preparations even without any metal; it indicated that the nerve and muscle were producing electricity by themselves. This finding laid the foundation of eleetro-physiology. The discovery that the messages in nervous system travel in the form of electric current replaced the neuroanatomical lesion technique by a less destructive and more precise neurophysiological technique for the study of behaviour. This technique involves two ways. 1. Recording of electrical activities from brain while animal is performing specific action oy ol nig 1 patterns, 2. Stimulating the brain region for eleciting behaviour.) 1. Recording of electric impulses. With the help of impulses in the brain are recorded and their correlation is f four basic types of brian waves have been recorded in hum () Alpha waves. These waves are recorded from the person is at rest with eyes closed, The brain is alert by Produced when we are awake but with a peaceful relaxed) (ii) Beta waves. These are generated from the sensory inputs. It means these waves indicate the active alert and is thinking or concentrating on some issue, (iii) Theta waves. These waves are produced from) emotional stress. They are associated with earliest stages — ~ 4 Animal Behaviow, Atphal WA Wy YI yy reel yaya Mn SVL yy 1 See] Fig. 2.6. Types of electric waves generated in different parts of the human brain. (iv) Delta waves. These waves are produced during deep sleep. 2. Initiation of behavioural patterns by electric stimulation. David Ferrier (1873) and Bartholow (1874) identified that particular motor patterns of muscular movement can be elicited by stimulating defined regions of the cerebral cortex of human brain. At the end of the 19th century, Ferrier was able to roughly localise the sensory centres in the cerebral cortex by vision, hearing (auditiory), taste (gustatory) and smell (olfactory). Electric stimulation method is the study of behavioural patterns that are initiated by electrical stimulation in specific brain regions by implanted electrodes. For example, J. Flynn (1929) stimulated a particular locus in the diencephalon of a cat and elicited prey catching behaviour. A. Zukerman (1965) was able to induce behaviours such as fleeing, courtship and threat by electrically stimulating different regions of a pigeon’s brain. E. von Hoist and Von Saint Paul (1960) elicited sleeping behaviour in a cock by stimulating the brain stem, By using'the same technique of electric stimulation by electrodes pleasure centres in human brainihave been identified by Jacobson (1967). 3. Neurochemical technique échniqu Nes! local application of drugs, i . eee neurotransmitters, some blo¢l Selectively. It is also called chemical brain stimu ee and bhang, istrated viour can be sterone) into Approaches to the Study of Behaviour 35 BOX 2.1. different ways. Depending upon their nature and been classified as follows: _ 1. Physiological agents. Adrenalin, histamine, serotonin and glutamic acid are known as physiological agents, They are present normally in the body but they are considered drugs when administered in concentrated dose. 2. Foreign agents. Reserpine, barbiturates and lysergic acid fall under this category. They are not normally present in the body. The chemicals ean affect behaviour chemical composition they b ‘Neuroactive drugs are also identified as tranquilizers, energizers or hypnotizers. Tranquilizers reduce agitation, excitement and emotion; energizers are stimulants such as caffine, amphetamine, ete. They tend to produce hyperexcitability and also facilitate sensory and motor perfomance. They also act as anti-depressants. Hypnotizers such as barbiturats are the drugs that produce sensory and motor depression and decp sleep. In large doses they cause unconciousness and even eat! hypothalamus. R.P. Mitchell (1968) found that administering the estrogen in anterior hypothalamus of brain of female cats whennot in heat produced mating behaviour. On the contrary, if the same hormone was injected in some other parts of brain, similar behaviour was produced. Most recently the brain is being, explored for its various functions using PET (i.e., Positron Emission Tomography) technique. 2.4. CAPTURING, MARKING AND TRACKING ANIMALS AND ANIMAL SIGNS Before discussing specific methods used for data collection in field settings, let us depart a moment to examine several related problems— issues that generally arise when we are conducting field studies, but that may also apply to some laboratory work in animal behaviour. These issues concern the methods of capturing animals, ways of marking animals for individual animals, and signs left by animals that may assist us with our interpretation of their behaviour. 1, Capturing Animals To study animals properly, it is often necessary to capture them. We may require to mark the animals for identification, We may require to make measurements of their size, sex or age. Or, we may wish to bring them into the laboratory for certain portion of our research work. ‘There are a variety of methods of capturing the animals that we can employ (Eltringham, 1978). Exact capture technique we use in a particular situation will depend upon the species and the nature of its habitat, The most common methods involved nets or traps. For many insect species we would use either sweep nets (Fig. 2.8) or dip nets. For birds we often use mist nest, nets of fine mesh set up using two poles, some what like a volleyball net. Mist nets are often set up to intersect the flight of the birds, for example, across a clearing in the woods or along the path of a stream. Larger nets can be used with small cannon to capture flocks of birds at a feeding station or mammals baited to come to a particular site. Live traps for vertebrates and invertebrates are available in an enormous variety of shapes and forms and can be used in the capture of almost all species of animals. Seine nets (Fig. 2.7) are often used to obtain samples of fish populations from a stream or lake. Pitfall traps can be placed into ground so their top rims are flush with the ground surface, are used for capturing a variety of invertebrates, reptiles, amphibians and mammals. Insects can be removed from tree trunks via the use of brushes or small vacuums (eg., Bulb aspi id they can be remoyed from vegetation by shaking or beating the plant ——=y 36 . A ial Behav, ae line Lead line Bag Running ing a Rope (=warp) Tractor Reel f@) The brail of stout pole sp —"~ Mechanized ip seine; B-Ring or Danish seine; C- Purse seine. Approaches to the Study of Behaviour 37 BOX 2.2. Sweeping and beating are most productive methods of collecting large number of insects. 1, Sweeping, The purpose of sweeping is to dislodge insects from the vegetation by means of an insect net, e.g.. sweep het, sweep net (Fig. 2.8) contains a stout and short handle attached Plankton, net dip not Fig. 2.8. Sweep nets, to a metallic ring having bag of muslin or canvas. The collector sweeps the net back and forth though the grass or bushes as he walks. During use, the contents of net should be frequently checked and the specimens sorted and placed in a proper vial. 2. Beating. The principle of beating is to hit a branch of a tree or shrub hard enough with a heavy stick in such a way that the insects and other arthropods fall on a tray or white sheet placed below from where they can be captured and collected. 3. Collection by aspirators. The aspirator is a simple suction apparatus used for collecting small fragile insects, mites and spiders. In bulb aspirator suctions is applied through the suction Fig.2.9, A bulb-aspirator. edi through the intake tube directly into the collecting via. can be used to stun large numbers of fish, which from their stunning, Finally, for many mammals, | 38 ‘Animal Behaviay particularly large beasts such as b tranquilizer dart. Animals shot in this manner are anesthetized for varying periods, and they are given ay antidote when the investigator has finished tagging or measuring them, 5 of ungulates, we employ a CO, cartridge gun ang, 2. Animal Identification During animal behaviour research, we want to be able to identify individual animals, Ouy rescarch questions might include: which animals are dominant ? Which females are mating with which males ? Are young animals cared for only by their mother, or do other members of group share in this process ? We also may often wish to avoid measuring the same animals twice by mistake, Folllwing techniques are used for animal-marking: Identification of animals of their natural marks. We can primarily identify animals by their natural marks (Pennycuik, 1978). Examples of the use of this technique include coat colour patterns in zebras (Equus burchelli) (Petersen, 1972) and giraffe (Giraffa camelopardalis) (Foster, 1966); differences in bill patterns in Bewick’s swans (Cygnus Columbianus) (Scott, 1978) and variation in physical charcteristics, natural mutilations and scars in primates (Ingram, 1978), lions (Panthera leo) (Pennycuik and Rudnai, 1970), and bottlenose dolphins (Wursig and Wursig, 1977). 2, Tagging. This technique for individual identification involves tagging animals; the nature of such tags will vary with the species (Stonehouse, 1978). Among the various tagging techniques used are metal and plastic leg bands in birds (Spencer, 1978; Patterson, 1978), fin clips or punches and metal fingerling tags for fish (Laired, 1978); toe clips, ear punches, dye marks, and tatoos for a variety of mammal species (Lane-Petter, 1978); and small dots of paint or dye on invertebrates (Southwood, 1994). Another animal-marking technique involves radioisotopes, cither in collars or bands of in subcutaneous implants. Because of potential problems with radioactivity and possible effects both on the animal bearing the radioisotope label and on the environment, the use of this technique has been curtailed considerably. More recently, Sheridan and Tamarin (1988) have developed the use of radio-nuclides for identification of individual meadow voles (Microtus pennsylvanicus) in order to study longevity and reproductive success; radionuclides are safer and not harmful to the animal carrying the label (see Drickamer ef al., 2002). We can also examine the DNA of animals, using several techniques; the most widely used is DNA fingerprinting. DNA can be extracted from blood or tissue samples or, simply, from hair or faeces. r 3. Tracking animals. Having marked animals for individual identification, we now might = ask: How can the animal be followed over time to study its behaviour? The most populat U technique of tracking the animals is radiotelemetry. An animal is captured and fitted with a t collar containing a radiotransmitter and battery or implanted with a similar device. Once released, ¢ the transmitter will send out signals that can be detected by a radio receiver and antenna. This a technology has been adapted for use with animals ranging in size from small birds, lizards and ' rodents, to whales and elephants, The size of the transmitter package and the range over which 1 he ick up the signal vary with the size of the animal and the requirement of the t wing an animal for many days, it is possible to obtain an accurate picture ‘ Uses and the portions of the day when it is active. Radio signals can now be ite circling the carth (Fig. 2.10); information from the satellite is then fed to i Olind station and animal movements can be analysed. This technique has t Y with caribou (Rangifer tarandus) in Alaska (Miller et al,, 1975) and ' 's turtles (see Mohanty, 2009), t ticularly large mammals or birds in flight, we may use vehicles t Approaches to the Study of Behaviour 39 aquatic animals, including man; a \y vertebrates (especially fish and whales) and a variety of invertebrates, we can use boats and diving gear to observe both from the water's surface and from below. In all of these situations, photographs or videotapes may aid in gathering information and enhance the process of identifying the individual and scoring the behaviour patterns, 4 Fig, 2.10, Satellite telemetry system, Shake-and-Bake Method. This technique has been developed for studying rodent movement by Kaufman in 1989. The procedure, somtimes called the “Shake-and -Bake” method, involves using a finely powdered fluorescent dye that comes in a variety of colours (e.g., magenta, lime green, blue, etc.). The animal to be tracked is captured and placed into a plastic bag with the powder. After gently shaking the bag for a few seconds, the pelage (viz., the coat or covering of a mammal, as of fur, wool, etc.) of the animal becomes covered with the dye, Once the animal is released, it will deposit a trail of dye for four to eight hours wherever it goes. On the night after the animal has been allowed to put down the dye trail, we go out with a special black light that causes light to fluoresce, and the trail of the animal can be followed quite easily. In this way, we obtain a record of the travels of the animal; the technique can be used with facility on both nocturnal and diurnal animals, 4. Animal signs. In fact, any trace, sign or construction which animals leave, can help us interpreting their behaviour. Examples of animal signs that we might expect to find include tracks, faecal material, egg shell, and animal remains. Almost all animals have tracks of some atic habitats, Impressions of feet can be found in mud, sand or other tracks to determing the direction of movement of an individual or of a group, and depending upon whether there are age or sex 40 ‘Animal Bohaviou, ditterences in the nature of the tracks, something about the age and sex of the individuay, 8? Examination of fecal material can tell us something of the diet of an animal. Ege, shells oe Felplus identify nest sites and indicate which species are nesting in a particular habit mo ve pains of animals might hold clues to the cause of death or could even tell the tale of an ia predation, Other, possibly more active signs we might discover would be nest, egg cases auf Prider webs. Nests of birds can provide important information about the site selection am x ae retruction of the nest, and when eggs or young are present, about the development of nes, ei Sampling Methods b Workers of animals behaviour use following sampling techniques: 1. Focal animal sampling. In recent years, field studies of many animal species haves focused much attention on research stractegies and methods of collecting data under felg cvaditions, Focal animal sampling (Altmann, 1974) involves recording all of the actions ang € jateractions of one particular animal during a prescribed time period. Using this method, ay “ abserver may watch a large number of animals, recording the behaviour of each For a shot period (e.., five or ten minutes per animals), or the observer may record the behaviour of veer animals, each being watched over a longer time period (hours per animal). As an example of the use of this technique we can consider here the study conducted by Bercovitch (1986) on Jominance rank and mating activity in male olive baboons (Papio anubis). The question tested concerning the relationship between male dominance rank and access (6 sexually receptive fomales for mating. Intense observations were made on only one or two baboons each day. The females that were the subjects or the focal sampling were in behavioural estrvs, exhibiting consort (companion) relationships with males. Using focal sampling, it was possible to record sll of the male partners of the sexually receptive females. When the data were analysed using Jnformation from only adult males that interacted with the receptive females, there was no clear miatioship between dominace status and mating activity. However, if adolescent males were Facluded in the analysis, then a relationship emerged, with dominant males engaging in more sae nal activity ith receptive females. One advantage of this observation technique is that intense samples can be gathered with a focus on particular animals. 2. Scan sampling or Instantaneous sampling. An observer using this technique watches each animal for only a few seconds at periodic intervals and records the activity (ies) that the nimal is performing only at the specific time marks indicated by the sampling scheme, The intervals between samples of the behaviour of each individual can vary, but generally, they a i few minutes to a half hour: As an example of the use of scan sampling method, consider # study of feeding behaviour of two captive groups of lemurs (Ganzhorn, 1986): the ring-tailed Jemur (Lemur catta) and the ruffed lemur (Varecia variegata) housed in seminatural environment Scan sampling was conducted on the groups of lemurs, using a thirty-minute interval betwee? records of individual activity. The species and part of the plant being eaten were recorded when we feu wae eating — at the time the data were to be recorded. By recording the data in this ae oe beerver pas ae to make many records on all the lemurs in both groups during all oad he analyses of these feeding data indicated that roughed lemurs spent les g the day, had longer feeding bouts, and consumed more fruits than ringtails BOX 2.3. out. The term bout is generally applied to (i) a repetitive occurrence of the same behaviour (e.g., about of pecking) or (i iv eee oa (ii)a relatively stereotyped sequence of behaviours that occur in a burst Approaches to the Study of Behaviour a One advantage of this technique is that it allows the observer to sample widely across all animals in the group and across a wide range of behaviour patterns, 3. One-zero sampling. It involves recording the occurrence or nonoccurrence of each of set of behaviour patterns within a series of time periods (Renner and Rosenzweig, 1986). This scheme is scen by some investigators as the best way to record a wide range of activities encompassing solitary actions, object-directed behaviour, and social interactions. This may also be a useful method for capturing the occurrence of behaviour patterns that either occur with very low frequency or are of brief duration. 4, Sequence sampling. This technique was used by Altamann (1974). In sequence sampling the focus is on the chain of behaviours. These may be performed by a single individual, eg. courtship displays in male ducks, parturition (viz, action or process of giving birth 0 offspring) in female, courtship display in stickleback fishes. The initiation of sample period is usually determined by the beginning of sequence and the sample period terminates where the observed sequence terminates. Some other sampling technique ad libitum (viz., without restraint or imposed | much or as often as is wanted by the animal) and all occurrence sampling, OF as 2.5. ROLE OF STATISTICS IN THE STUDY OF BEHAVIOUR Statistical data regarding different aspects of behaviour is important in deriving conclusions from various experiments. It is used to summarise and analyze data and to describe variability, differences and similarities and to determine the appropriate taxonomic classification of ambiguous species and classification of whole groups of species 1. Discrimination of different species. The animal species that are morphologically so similar that they can not be discriminated are defined on the basis oftheir breeding patterns and reproductive isolation. For example, a number of species of fireflies are identified by their Characteristic courtship pattern and they do not interbreed. This judgement is possible only by ‘maintaining their statistical records. 2. Classification of ambiguous species. Data from animal behaviour (ethology) can be used in determining appropriate taxonomic classification for ambiguous species and determining thelr similarities and derivation (ancestry). Bekoff, Hill and Milton (1975) determined the proper taxonomic position of New England canid relatives in relation to other canids by using cep vioural data, It supported the anatomical data that New England canids are more closely related to coyotes then to wolves. 3, Classification of whole groups of species. Behavioural data is also useful in the calssification of groups. For example, different species of ducks and geese display different behavioural patterns. These have been used in revising their taxonomic classification. ‘Analysis of variation and variance. Animals exhibit variation in appearance of performance for most morphological, physiological and behavioural traits. For example, some blackbirds (An American bird, Agelaius phoeniceus) perform territorial eatedly in the face of an intruder attempting to enter the territory. The ieoften associated with the conkaree (conch-like) call. While others display ider the same circumstances. Similarly, nests constructed by redwings bove|ground and the types of plant material used in construction. How Rental procedures to account for this variation? ter discrete or continuous. Diserete variation involves those measures ain values; For example, clutch size (number of eggs layed in a nest) sure; a species may lay 1, 2, 3 or upto 6 or more eggs per clutch, but

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