Behaviour

Tinbergen, N.

doi: 10.1163/156853960X00098pmid: N/A

<jats:sec><jats:title>Abstract</jats:title><jats:p>This paper describes a number of displays of various gulls (Laridae) with special reference to the Herring Gull group, the hooded gulls, and the Kittiwake, and discusses their functions, causation, evolutionary origin and further evolution as signals. After a sketch of the rationale of comparative behaviour studies (chapter II), the most common single displays and display sequences are described (chapter III). They are rather similar through the family, though many species-specific differences exist. The behaviour similarities strengthen the conclusion that the gulls are a monophyletic group. Evidence is presented in chapter IV to show that the displays have signal function. A distinction is made between distance-increasing and distance-reducing displays. The reasons for the occurence of more than one, yet no more than about six distance- increasing postures are discussed ; it is argued that each display may well be adapted to deal with a distinct category of opponent : accidental trespassers require another repellent than intentional persistent intruders; and within each category actual trespassers are met in another way than potential trespassers. In addition, one particularly loud call is a typical long-distance threat. This call usually acts as advertisement in a double sense: it repels competitors and attracts unmated females. Some of the distance-reducing (or appeasement) postures are used in agonistic situations, and still more regularly at the end of the pair formation or meeting ceremony. The need for and the possibility of more systematic studies of the precise functions of displays is stressed. Chapter V discusses aims and methods of analysis of the motivation of the displays. The value of "natural experiments" is stressed. The application of three independent methods leads to the conclusion that agonistic displays are ambivalent, i.e. the outcome of the simultaneous arousal of a tendency to attack and a tendency to flee: the relation between fluctuations in these tendencies and the displays shown is examined. There is a striking correlation between the motivation of these displays and the information they pass on to other individuals. Appeasement gestures always contain an element of fear; this tendency is in conflict with a tendency to stay, which can be, but usually is not part of the tendency to attack; it may be sexual attraction, attraction to at nest site, or attraction to a provider of food. The need for motivational analyses of the many different forms in which one posture can occur is stressed. The similarity of motivation in the pair formation ceremonies in the diff.erent species is much greater than the formal similarity of the display sequences; the conflict theory of courtship is tested. The origin of the displays (chapter VI) is varied. Some have clearly arisen as preparatory or intention movements of the patterns directly aroused by the situation ("aut- ochthonous" movements) ; of these, some are redirected to inanimate objects. Others are derived from movements belonging to functional patterns not directly aroused by the situation ("displacement activities") ; their various origins are discussed, and it is shown that they are second components of a dual movement, of which the first component is "auto?hthonous" in the above sense and facilitates the displacement activity. In chapter VII some ultimate causes of evolutionary change are discussed, and a preliminary functional classification of alleged changes in displays is presented. It is argued that change has been enhanced by at least four different types of selection pressure: (i) towards improvement of signal function (conspicuousness) ; (2) towards increase in intraspecific unambiguity; (3) towards increased interspecific unambiguity; and (4) as a corrollary of selection pressure in other functional systems. In some gulls, (4) may have made the major contribution to sexual isolation. A preliminary classification is given of the postulated evolutionary changes in behaviour mechanisms. The value of behaviour characters for taxonomic use is considered. After a discussion of adaptive and non-adaptive differences the relative validity of LORLNz's emphasis on the phylogenetic conservatism of displays is reconsidered, and the occurrence of convergent similarities is demonstrated.</jats:p> </jats:sec>

Tinbergen, N.

doi: 10.1163/156853960x00098pmid: N/A

AbstractThis paper describes a number of displays of various gulls (Laridae) with special reference to the Herring Gull group, the hooded gulls, and the Kittiwake, and discusses their functions, causation, evolutionary origin and further evolution as signals. After a sketch of the rationale of comparative behaviour studies (chapter II), the most common single displays and display sequences are described (chapter III). They are rather similar through the family, though many species-specific differences exist. The behaviour similarities strengthen the conclusion that the gulls are a monophyletic group. Evidence is presented in chapter IV to show that the displays have signal function. A distinction is made between distance-increasing and distance-reducing displays. The reasons for the occurence of more than one, yet no more than about six distance- increasing postures are discussed ; it is argued that each display may well be adapted to deal with a distinct category of opponent : accidental trespassers require another repellent than intentional persistent intruders; and within each category actual trespassers are met in another way than potential trespassers. In addition, one particularly loud call is a typical long-distance threat. This call usually acts as advertisement in a double sense: it repels competitors and attracts unmated females. Some of the distance-reducing (or appeasement) postures are used in agonistic situations, and still more regularly at the end of the pair formation or meeting ceremony. The need for and the possibility of more systematic studies of the precise functions of displays is stressed. Chapter V discusses aims and methods of analysis of the motivation of the displays. The value of "natural experiments" is stressed. The application of three independent methods leads to the conclusion that agonistic displays are ambivalent, i.e. the outcome of the simultaneous arousal of a tendency to attack and a tendency to flee: the relation between fluctuations in these tendencies and the displays shown is examined. There is a striking correlation between the motivation of these displays and the information they pass on to other individuals. Appeasement gestures always contain an element of fear; this tendency is in conflict with a tendency to stay, which can be, but usually is not part of the tendency to attack; it may be sexual attraction, attraction to at nest site, or attraction to a provider of food. The need for motivational analyses of the many different forms in which one posture can occur is stressed. The similarity of motivation in the pair formation ceremonies in the diff.erent species is much greater than the formal similarity of the display sequences; the conflict theory of courtship is tested. The origin of the displays (chapter VI) is varied. Some have clearly arisen as preparatory or intention movements of the patterns directly aroused by the situation ("aut- ochthonous" movements) ; of these, some are redirected to inanimate objects. Others are derived from movements belonging to functional patterns not directly aroused by the situation ("displacement activities") ; their various origins are discussed, and it is shown that they are second components of a dual movement, of which the first component is "auto?hthonous" in the above sense and facilitates the displacement activity. In chapter VII some ultimate causes of evolutionary change are discussed, and a preliminary functional classification of alleged changes in displays is presented. It is argued that change has been enhanced by at least four different types of selection pressure: (i) towards improvement of signal function (conspicuousness) ; (2) towards increase in intraspecific unambiguity; (3) towards increased interspecific unambiguity; and (4) as a corrollary of selection pressure in other functional systems. In some gulls, (4) may have made the major contribution to sexual isolation. A preliminary classification is given of the postulated evolutionary changes in behaviour mechanisms. The value of behaviour characters for taxonomic use is considered. After a discussion of adaptive and non-adaptive differences the relative validity of LORLNz's emphasis on the phylogenetic conservatism of displays is reconsidered, and the occurrence of convergent similarities is demonstrated.

Sawin, Paul B.; Petropolus, Stephen F.; Denenberg, Victor H.; Ross, Sherman

doi: 10.1163/156853960X00106pmid: N/A

<jats:sec><jats:title>Abstract</jats:title><jats:p>The records of a number of female rabbits of races III, IIIc, X, and AcEp which had produced at least four litters were analyzed for correlations between scattering and cannibalism and five other measures of maternal behavior. These rabbits were classed as low newborn mortality mothers and the maternal behavior scores for their first successful litter were compared with the behavioral scores of high newborn mortality mothers (60 per cent mortality of complete litters) for their first successful litter. Only race IIIc showed any significant amount of scattering and cannibalism. In this race, correlational analyses of scattering and cannibalism against the five other measures of maternal behavior found that both were significantly related to nest quality and had weaker, though positive, correlations with time of nest building. Race IIIc does were only moderately consistent in their scattering or cannibalizing behavior over four litters and these variables were correlated .404. No differences were found between high and low newborn mortality mothers except for race AcEp, where the low mortality mothers were found to have significantly more interest in their young and to be better nurse-mothers than the high newborn mortality does. The findings from this study and a previous one (DENENBERG, et al., 1958) indicate that better nest quality, earlier time of nest building, greater percentage of live born young suckled on the first day, lack of scattering, and lack of cannibalism are inter-correlated. It was concluded that these characteristics can be classified as a "maternal care" complex which appears to be independent of a second group involving interest in young and aggressive protection.</jats:p> </jats:sec>

Denenberg, Victor H.; Petropolus, Stephen F.; Sawin, Paul B.; Ross, Sherman

doi: 10.1163/156853960x00106pmid: N/A

AbstractThe records of a number of female rabbits of races III, IIIc, X, and AcEp which had produced at least four litters were analyzed for correlations between scattering and cannibalism and five other measures of maternal behavior. These rabbits were classed as low newborn mortality mothers and the maternal behavior scores for their first successful litter were compared with the behavioral scores of high newborn mortality mothers (60 per cent mortality of complete litters) for their first successful litter. Only race IIIc showed any significant amount of scattering and cannibalism. In this race, correlational analyses of scattering and cannibalism against the five other measures of maternal behavior found that both were significantly related to nest quality and had weaker, though positive, correlations with time of nest building. Race IIIc does were only moderately consistent in their scattering or cannibalizing behavior over four litters and these variables were correlated .404. No differences were found between high and low newborn mortality mothers except for race AcEp, where the low mortality mothers were found to have significantly more interest in their young and to be better nurse-mothers than the high newborn mortality does. The findings from this study and a previous one (DENENBERG, et al., 1958) indicate that better nest quality, earlier time of nest building, greater percentage of live born young suckled on the first day, lack of scattering, and lack of cannibalism are inter-correlated. It was concluded that these characteristics can be classified as a "maternal care" complex which appears to be independent of a second group involving interest in young and aggressive protection.

Franzisket, L.

doi: 10.1163/156853960X00115pmid: N/A

<jats:sec><jats:title>Abstract</jats:title><jats:p>Durch Wahlversuche wird gezeigt, dass die auffällige Schwarz-Weiss-Färbung des Preussenfisches Dascyllus aruanus keine kryptische Wirkung, sondern attraktive Wirkung auf Artgenossen ausübt. Durch die Streifung sind die Fische zwar gegen ihre Umgebung auffällig, aber nicht als Beuteschema "Fischgestalt" auffällig.</jats:p> </jats:sec>

Franzisket, L.

doi: 10.1163/156853960x00115pmid: N/A

AbstractDurch Wahlversuche wird gezeigt, dass die auffällige Schwarz-Weiss-Färbung des Preussenfisches Dascyllus aruanus keine kryptische Wirkung, sondern attraktive Wirkung auf Artgenossen ausübt. Durch die Streifung sind die Fische zwar gegen ihre Umgebung auffällig, aber nicht als Beuteschema "Fischgestalt" auffällig.

Jensen, Donald D.

doi: 10.1163/156853960x00124pmid: N/A

AbstractParamecium aurelia is the subject of a theory of behavior composed of postulates describing the presence, action, and interaction of three pacemakers: a posterior pacemaker which produces ciliary beat backward along the kineties; a buccal pacemaker which produces ciliary beat toward the suture line and the buccal overture in the vestibulum ; and an anterior pacemaker which produces ciliary beat away from the suture line and the buccal overture in the vestibulum and forward along the kineties elsewhere. Commonly observed behaviors of paramecia are derived from the theory and six experiments relevant to the theory are described. In three experiments, the culture medium in which paramecia were observed was varied. Presence of bacteria and non-nutritive particulate matter decreased swimming scores and increased swimming turn rates. In one of these experiments prior as well as immediate presence of bacteria in culture fluid varied. Prior availability of food bacteria increased swimming scores and contact turn rates. These results are consistent with the theory if it is assumed that immediate presence of particulate matter increases buccal pacemaker activity and that prior availability of food bacteria increases posterior pacemaker activity. In another experiment animals were observed during and following binary fission. During fission swimming scores decreased, and following fission the proters had lower swimming scores and contact turn rates than opisthes. These results are consistent with the theory if it is assumed that periods in which all ciliary beat is absent occur in parent animals during fission, and in proters following fission. Such periods were observed in dividing animals. Opisthes had higher swimming turn rates following fission than prior to it; this result is consistent with the theory if it is assumed that newly organized buccal pacemakers are unusually active. In two other experiments, the anterior tips, buccal cavities, and posterior tips of paramecia were subjected to ultraviolet microbeam irradiation. Irradiated animals moved more during and immediately following irradiation than they moved prior to irradiation or than control animals did. Posteriorly irradiated animals moved forward, anteriorly irradiated animals moved backward. Irradiation also influenced behavior during a subsequent two-minute observation period. Anteriorly irradiated animals had higher swimming scores, lower contact turn rates, and a greater tendency to swim backward. Animals irradiated in the buccal cavity had lower swimming turn rates and higher swimming turn durations; posteriorly irradiated animals had lower swimming scores, contact turn rates, and swimming turn rates. These results are consistent with the theory if pacemakers near irradiated areas are temporarily activated by irradiation and are hyposensitive and hyperreactive following irradiation. A number of research problems were suggested.

Jensen, Donald D.

doi: 10.1163/156853960X00124pmid: N/A

<jats:sec><jats:title>Abstract</jats:title><jats:p>Paramecium aurelia is the subject of a theory of behavior composed of postulates describing the presence, action, and interaction of three pacemakers: a posterior pacemaker which produces ciliary beat backward along the kineties; a buccal pacemaker which produces ciliary beat toward the suture line and the buccal overture in the vestibulum ; and an anterior pacemaker which produces ciliary beat away from the suture line and the buccal overture in the vestibulum and forward along the kineties elsewhere. Commonly observed behaviors of paramecia are derived from the theory and six experiments relevant to the theory are described. In three experiments, the culture medium in which paramecia were observed was varied. Presence of bacteria and non-nutritive particulate matter decreased swimming scores and increased swimming turn rates. In one of these experiments prior as well as immediate presence of bacteria in culture fluid varied. Prior availability of food bacteria increased swimming scores and contact turn rates. These results are consistent with the theory if it is assumed that immediate presence of particulate matter increases buccal pacemaker activity and that prior availability of food bacteria increases posterior pacemaker activity. In another experiment animals were observed during and following binary fission. During fission swimming scores decreased, and following fission the proters had lower swimming scores and contact turn rates than opisthes. These results are consistent with the theory if it is assumed that periods in which all ciliary beat is absent occur in parent animals during fission, and in proters following fission. Such periods were observed in dividing animals. Opisthes had higher swimming turn rates following fission than prior to it; this result is consistent with the theory if it is assumed that newly organized buccal pacemakers are unusually active. In two other experiments, the anterior tips, buccal cavities, and posterior tips of paramecia were subjected to ultraviolet microbeam irradiation. Irradiated animals moved more during and immediately following irradiation than they moved prior to irradiation or than control animals did. Posteriorly irradiated animals moved forward, anteriorly irradiated animals moved backward. Irradiation also influenced behavior during a subsequent two-minute observation period. Anteriorly irradiated animals had higher swimming scores, lower contact turn rates, and a greater tendency to swim backward. Animals irradiated in the buccal cavity had lower swimming turn rates and higher swimming turn durations; posteriorly irradiated animals had lower swimming scores, contact turn rates, and swimming turn rates. These results are consistent with the theory if pacemakers near irradiated areas are temporarily activated by irradiation and are hyposensitive and hyperreactive following irradiation. A number of research problems were suggested.</jats:p> </jats:sec>

Manning, Aubrey

doi: 10.1163/156853960x00133pmid: N/A

Abstract1. The courtship behaviour of Drosophila melanogaster and D. simulans has been studied. Melanogaster is a generally more active species than simulans and its courtship is more vigorous. 2. The sexual behaviour of both types of male can be readily classified into the same basic elements, orientation, wing display, licking and attempted copulation. 3. Simulans males have longer lag periods before courtship, longer bouts of simple orientation, fewer changes of element when courting and their chief wing display is 'scissoring'. Melanogaster's chief wing display is 'vibration'. 4. Experiments in which the sexual stimulus situation was changed show that with lower stimulation melanogaster males show an increased proportion of scissoring, while with increased stimulation simulans males show more vibration. Additional reasons are given for considering scissoring to represent a lower level of sexual excitation than vibration. 5. The differences between the males can be most simply ascribed to simulans having a slower rise of sexual excitation during courtship than melanogaster, not to any changes in the basic organization of their behaviour. This slowed rise may be related to higher thresholds of response in simulans. 6. The females of simulans are more responsive to visual aspects of their males' courtship, and less responsive to those stimuli perceived by their antennae, than are melanogaster females. 7. It is argued that these changes in courtship behaviour probably resulted from selection acting upon a common ancestral population so as to favour high activity in some environments and low activity in others. These changes will be most readily produced by mutations affecting reaction thresholds and the changes to courtship pattern will follow. Selection will then favour females responsive to the most easily available stimuli from the males' courtship. 8. It is suggested that similar situations occur in other sibling Drosophila species and that as a first step in behavioural divergence, mutations affecting thresholds will be particularly important.

Manning, Aubrey

doi: 10.1163/156853960X00133pmid: N/A

<jats:sec><jats:title>Abstract</jats:title><jats:p>1. The courtship behaviour of Drosophila melanogaster and D. simulans has been studied. Melanogaster is a generally more active species than simulans and its courtship is more vigorous. 2. The sexual behaviour of both types of male can be readily classified into the same basic elements, orientation, wing display, licking and attempted copulation. 3. Simulans males have longer lag periods before courtship, longer bouts of simple orientation, fewer changes of element when courting and their chief wing display is 'scissoring'. Melanogaster's chief wing display is 'vibration'. 4. Experiments in which the sexual stimulus situation was changed show that with lower stimulation melanogaster males show an increased proportion of scissoring, while with increased stimulation simulans males show more vibration. Additional reasons are given for considering scissoring to represent a lower level of sexual excitation than vibration. 5. The differences between the males can be most simply ascribed to simulans having a slower rise of sexual excitation during courtship than melanogaster, not to any changes in the basic organization of their behaviour. This slowed rise may be related to higher thresholds of response in simulans. 6. The females of simulans are more responsive to visual aspects of their males' courtship, and less responsive to those stimuli perceived by their antennae, than are melanogaster females. 7. It is argued that these changes in courtship behaviour probably resulted from selection acting upon a common ancestral population so as to favour high activity in some environments and low activity in others. These changes will be most readily produced by mutations affecting reaction thresholds and the changes to courtship pattern will follow. Selection will then favour females responsive to the most easily available stimuli from the males' courtship. 8. It is suggested that similar situations occur in other sibling Drosophila species and that as a first step in behavioural divergence, mutations affecting thresholds will be particularly important.</jats:p> </jats:sec>