Distant Food Sources Research Articles

Ants (Lasius niger) deposit more pheromone close to food sources and further from the nest but do not attempt to update erroneous pheromone trails

Many social insects use pheromone trails to recruit workers to resources. Pheromone trails have some limitations, however. Two major limitations are that they evaporate over time, meaning that more distant locations may be more difficult to recruit precisely to, and that they cannot be removed if laid erroneously, or if they become outdated. Here, we ask whether Lasius niger ants can adaptively modulate pheromone deposition to overcome these limitations. Specifically, we first ask whether ants which have followed a pheromone trail to a wrong (unrewarded) location upregulate pheromone deposition to the correct food location to compete with the erroneous trail. They do not. Then, we examine how food distance and proximity to food influence pheromone deposition. We find that ants deposit up to 22 times more pheromone within 10 cm of a food source compared to when they are about to reach the nest. Moreover, they deposit up to four times more pheromone next to a food source placed 100 cm away from the nest as compared to one placed 20 cm away from the nest. Lasius niger ants thus do not display a mechanism for overcoming outdated or erroneously-laid trails by strengthening correct alternatives in our experiment, but do strongly upregulate recruitment to more distant food sources, which are more difficult to locate and recruit to.

The dive performance of immature king penguins following their annual molt suggests physiological constraints.

Like all birds, penguins undergo periodic molt, during which they replace old feathers. However, unlike other birds, penguins replace their entire plumage within a short period while fasting ashore. During molt, king penguins (Aptenodytes patagonicus) lose half of their initial body mass, most importantly their insulating subcutaneous fat and half of their pectoral muscle mass. The latter might challenge their capacity to generate and sustain a sufficient mechanical power output to swim to distant food sources and propel themselves to great depth for successful prey capture. To investigate the effects of the annual molt fast on their dive/foraging performance, we studied various dive/foraging parameters and peripheral temperature patterns in immature king penguins across two molt cycles, after birds had spent their first and second year at sea, using implanted data-loggers. We found that the dive/foraging performance of immature king penguins was significantly reduced during post-molt foraging trips. Dive and bottom duration for a given depth were shorter during post-molt and post-dive surface interval duration was longer, reducing overall dive efficiency and underwater foraging time. We attribute this decline to the severe physiological changes that birds undergo during their annual molt. Peripheral temperature patterns differed greatly between pre- and post-molt trips, indicating the loss of the insulating subcutaneous fat layer during molt. Peripheral perfusion, as inferred from peripheral temperature, was restricted to short periods at night during pre-molt but occurred throughout extended periods during post-molt, reflecting the need to rapidly deposit an insulating fat layer during the latter period.

Ants (Temnothorax nylanderi) adjust tandem running when food source distance exposes them to greater risks

Social insect colonies exploit food sources that vary in their profitability and riskiness. One factor that affects both profitability and riskiness is the foraging distance: more distant resources are both more costly to exploit and expose individuals to greater predation or navigational risks. Temnothorax nylanderi scouts use tandem running to recruit nestmates to resources, such as food or nest sites. Tandem runs are often unsuccessful, leaving followers in potentially unknown or dangerous territory. Thus, as foraging distances increase, communication mistakes are likely to be more costly. We tested if leaders and followers adjust their tandem running behavior in response to increasing foraging distances. We asked whether the success rate, the probability to recruit, and the waiting time following a loss of contact depend on the foraging distances. We found that the success rate (75–86%) of tandem runs does not decrease with increasing foraging distance but rapidly increases with the leader’s experience, from 67% for the first tandem run to 94% for the fourth. Pairs progressed faster, and followers search longer for their partner after a loss of contact when visiting more distant food sources. The probability to perform a tandem run did not decrease with the foraging distance but increased with foraging experience. Our results indicate that ants might attempt to reduce exposure to risks by progressing faster when visiting more distant food sources. As ants become more experienced, they lead more and better tandem runs. These findings suggest that both leaders and followers respond to the potential dangers posed by exploiting faraway resources. Foraging distance plays an important ecological role in animals as the foraging distance affects both energetic costs and predation risk. Ants have evolved several cooperative foraging strategies to exploit a food source as efficiently as possible, including a recruitment method called “tandem running.” Here, an informed leader guides a naive follower to a valuable resource. We tested if tandem running behavior changes if food sources are more distant. Foraging distance indeed had an impact as leaders walked faster and followers searched for longer after a contact loss when food sources were more distant.

Tandem Recruitment and Foraging in the Ponerine Ant Pachycondyla harpax (Fabricius).

Tandem running is a common recruitment strategy in ant species with small colony sizes. During a tandem run, an informed leader guides a usually naïve nestmate to a food source or a nest site. Some species perform tandem runs only during house hunting, suggesting that tandem running does not always improve foraging success in species known to use tandem running as a recruitment strategy, but more natural history information on tandem running under natural conditions is needed to better understand the adaptive significance of tandem recruitment in foraging. Studying wild colonies in Brazil, we for the first time describe tandem running in the ponerine ant Pachycondyla harpax (Fabricius). We asked if foragers perform tandem runs to carbohydrate- (honey) and protein-rich (cheese) food items. Furthermore, we tested whether the speed and success rate of tandem runs depend on the foraging distance. Foragers performed tandem runs to both carbohydrate food sources and protein-rich food items that exceed a certain size. The probability to perform a tandem run and the travelling speed increase with increasing foraging distances, which could help colonies monopolize more distant food sources in a competitive environment. Guiding a recruit to a food source is costly for leaders as ants are ~66% faster when travelling alone. If tandem runs break up (~23% of all tandem runs), followers do not usually discover the food source on their own but return to the nest. Our results show that tandem running to food sources is common in P. harpax, but that foragers modify their behaviour according to the type of food and its distance from the nest. Competition with other ants was intense and we discuss how tandem running in P. harpax might help colonies to build-up a critical number of ants at large food items that can then defend the food source against competitors.

Exploring the chemo-textural familiarity hypothesis for scorpion navigation

The navigation by scene familiarity hypothesis provides broad explanatory power for how bees and ants navigate from the hive to distant food sources and back. The premise is that the visual world is decomposed into pixelated matrices of information that are stored and readdressed as the insects retrace learned routes. Innate behaviors in these insects (including learning walks/flights and path integration) provide the important goal-directed views to allow the initial retracing (i.e., the insect must learn the scene while moving toward the goal because everything looks different while moving away). Scorpion navigation may use a similar premise, with the chemical and textural features of the environment substituting for visual input. Scorpion pectines support thousands of chemo- and mechano-sensitive units called peg sensilla, each containing at least 10 energetically expensive sensory neurons. We have long wondered why pectines have so many pegs and associated neurons. Many sand scorpions emerge onto the surface from their home burrows at night to pursue insect prey and somehow find their way back to their burrows. Based on the measured resolution of peg sensilla, we have calculated that sufficient information exists in sand's texture to enable scorpions to retrace previously experienced paths with little to no chance of confusion. Preliminary evidence of learning walks and path integration in scorpions is also congruent with the navigation by chemo-textural familiarity hypothesis.

Water Transport byCoptotermes formosanus(Isoptera: Rhinotermitidae)

Subterranean termites are extremely vulnerable to desiccation, and high moisture makes their habitat and food favorable for survival and colony growth. Although there is a general perception that termites can manipulate moisture, documentation is surprisingly scanty with regard to how termites transport water and the factors that impact it. There has been no study of water transfer by Coptotermes formosanus Shiraki, a notoriously invasive termite in the southern United States. We conducted a study to determine if C. formosanus transfers water. Bioassays using arenas with a dry food source connected to a moist substrate by either a short tube (10 cm) or a long tube (100 cm) were conducted. Three moistened substrate types were tested to see how they impacted water transfer. In addition, workers and soldiers sampled from a moist sand substrate were dissected to determine water sac volumes for possible transfer of water to wood. The results indicated that some water transfer is achieved by the evacuation of water sacs. However, moist soil was also moved to increase humidity. When termites had use of moist silty clay, wood moisture gain increased significantly in both 10 and 100 cm tubes. As tube distance increased, moisture to the more distant food source decreased. Workers had the largest water sacs, though soldiers appear to contribute in water transfer via water sacs as well. Water transfer and its implications are discussed.

Comparative population densities of three species of doves (Columbidae) in disturbed landscapes in Northern Paraná State, Brazil

Assessments of population density and contact number of individuals of the Eared Dove Zenaida auriculata were conducted in areas of the city of Londrina and were compared with those of two other species of the family Columbidae (Columbina talpacoti and C. picui). Data were collected from distance sampling in linear transects of 2,000 m. Analyses of density were calculated using the program DISTANCE. The results show that C. talpacoti and C. picui were recorded in all months in urban areas, and less frequently in rural and pasture areas. Zenaida auriculata was the most frequently recorded species in all areas, showing high rates of population density with seasonal variations; density peaks were observed during September and January in the urban area (coinciding with the arrival of warmer days in the region), while in rural areas, density peaks were also recorded during November/ December, March and April/May — periods in which planting of soybean and maize, and the soybean harvest, and wheat planting in the region, occurred, respectively. The density of Z. auriculata during these months showed a value that was 93 times greater in November/December and 75 times greater in April/May, compared to the density in the other months of the year. Although Z. auriculata was considered uncommon a few decades ago, we suggest that the removal of the original forest and the expansion of agriculture in the region have provided a large amount of seeds in the environment, which might favor the increased density of Z. auriculata in the region. The high rate of population density achieved by the Eared Dove suggests that the species has found a favorable environment in the region. This includes the ease of obtaining resources, suitable sites for nesting, reproductive success and possible adaptive factors, such as the opportunist movement in search of distant food sources.

Assessing sex-related chick provisioning in greater flamingo Phoenicopterus roseus parents using capture-recapture models.

In sexually dimorphic species, the parental effort of the smaller sex may be reduced due to competitive exclusion in the feeding areas by the larger sex or physiological constraints. However, to determine gender effects on provisioning patterns, other intrinsic and extrinsic factors affecting parental effort should be accounted for. Greater flamingos (Phoenicopterus roseus) exhibit sexual size dimorphism. In Fuente de Piedra colony, the lake dries out almost completely during the breeding season and both parents commute between breeding and foraging sites >130 km away during the chick-rearing period. Applying multistate capture-recapture models to daily observations of marked parents, we determined the effects of sex, and their interactions with other intrinsic and extrinsic factors, on the probability of chick desertion and sojourn in the colony and feeding areas. Moreover, using stable isotopes in the secretions that parents produce to feed their chicks, we evaluated sex-specific use of wetlands. The probability of chick attendance (complementary to chick desertion) was >0.98. Chick desertion was independent of parental sex, but decreased with parental age. Females stayed in the feeding areas for shorter periods [mean: 7.5 (95% CI: 6.0-9.4) days] than males [9.2 (7.3-11.8) days]. Isotopic signatures of secretions did not show sex differences in δ(13)C, but males' secretions were enriched in δ(15)N, suggesting they fed on prey of higher trophic levels than females. Both parents spent approximately 1 day in the colony, but females prolonged their mean stay when the lake dried out. Females also allocated more time to foraging in the flooded areas remaining in the colony, likely because they were energetically more stressed than males. The results indicate that sex-specific provisioning behaviour in greater flamingo is related to differential effects of both intrinsic and extrinsic factors. Males seem forage less efficiently than females, whereas females' body condition seems to be lower after feeding the chick. Our methodology may be extended to species that feed on distant food sources and that do not visit their offspring daily, to elucidate patterns of chick-provisioning behaviour.

Drosophila Tracks Carbon Dioxide in Flight

Carbon dioxide (CO(2)) elicits an attractive host-seeking response from mosquitos yet is innately aversive to Drosophila melanogaster despite being a plentiful byproduct of attractive fermenting food sources. Prior studies used walking flies exclusively, yet adults track distant food sources on the wing. Here we show that a fly tethered within a magnetic field allowing free rotation about the yaw axis actively seeks a narrow CO(2) plume during flight. Genetic disruption of the canonical CO(2)-sensing olfactory neurons does not alter in-flight attraction to CO(2); however, antennal ablation and genetic disruption of the Ir64a acid sensor do. Surprisingly, mutation of the obligate olfactory coreceptor (Orco) does not abolish CO(2) aversion during walking yet eliminates CO(2) tracking in flight. The biogenic amine octopamine regulates critical physiological processes during flight, and blocking synaptic output from octopamine neurons inverts the valence assigned to CO(2) and elicits an aversive response in flight. Combined, our results suggest that a novel Orco-mediated olfactory pathway that gains sensitivity to CO(2) in flight via changes in octopamine levels, along with Ir64a, quickly switches the valence of a key environmental stimulus in a behavioral-state-dependent manner.

Honey bee waggle dance communication: signal meaning and signal noise affect dance follower behaviour

Returning honey bee foragers perform waggle dances to inform nestmate foragers about the presence, location and odour of profitable food sources and new nest sites. The aim of this study is to investigate how the characteristics of waggle dances for natural food sources and environmental factors affect dance follower behaviour. Because food source profitability tends to decrease with increasing foraging distance, we hypothesised that the attractiveness of a dance, measured as the number of dance followers and their attendance, decreases with increasing distance to the advertised food location. Additionally, we determined whether time of year and dance signal noise, quantified as the variation in waggle run direction and duration, affect dance follower behaviour. Our results suggest that bees follow fewer waggle runs as the food source distance increases, but that they invest more time in following each dance. This is because waggle run duration increases with increasing foraging distance. Followers responded to increased angular noise in dances indicating more distant food sources by following more waggle runs per dance than when angular noise was low. The number of dance followers per dancing bee was also affected by the time of year and varied among colonies. Our results provide evidence that both noise in the message, that is variation in the direction component, and the message itself, that is the distance of the advertised food location, affect dance following. These results indicate that dance followers may pay attention to the costs and benefits associated with using dance information.

Assessing Burrowing, Nest Construction, and Hoarding in Mice

Deterioration in the ability to perform "Activities of daily living" (ADL) is an early sign of Alzheimer's disease (AD). Preclinical behavioural screening of possible treatments for AD currently largely focuses on cognitive testing, which frequently demands expensive equipment and lots of experimenter time. However, human episodic memory (the most severely affected aspect of memory in AD) is different to rodent memory, which seems to be largely non-episodic. Therefore the present ways of screening for new AD treatments for AD in rodents are intrinsically unlikely to succeed. A new approach to preclinical screening would be to characterise the ADL of mice. Fortuitously, several such assays have recently been developed at Oxford, and here the three most sensitive and well-characterised are presented. Burrowing was first developed in Oxford13. It evolved from a need to develop a mouse hoarding paradigm. Most published rodent hoarding paradigms required a distant food source to be linked to the home cage by a connecting passage. This would involve modifying the home cage as well as making a mouse-proof connecting passage and food source. So it was considered whether it would be possible to put the food source inside the cage. It was found that if a container was placed on the floor it was emptied by the next morning., The food pellets were, however, simply deposited in a heap at the container entrance, rather than placed in a discrete place away from the container, as might be expected if the mice were truly hoarding them. Close inspection showed that the mice were performing digging ("burrowing") movements, not carrying the pellets in their mouths to a selected place as they would if truly hoarding them.6 Food pellets are not an essential substrate for burrowing; mice will empty tubes filled with sand, gravel, even soiled bedding from their own cage. Moreover, they will empty a full tube even if an empty one is placed next to it8. Several nesting protocols exist in the literature. The present Oxford one simplifies the procedure and has a well-defined scoring system for nest quality5. A hoarding paradigm was later developed in which the mice, rather than hoarding back to the real home cage, were adapted to living in the "home base" of a hoarding apparatus. This home base was connected to a tube made of wire mesh, the distal end of which contained the food source. This arrangement proved to yield good hoarding behaviour, as long as the mice were adapted to living in the "home base" during the day and only allowed to enter the hoarding tube at night.

Assessing Burrowing, Nest Construction, and Hoarding in Mice

Deterioration in the ability to perform "Activities of daily living" (ADL) is an early sign of Alzheimer's disease (AD). Preclinical behavioural screening of possible treatments for AD currently largely focuses on cognitive testing, which frequently demands expensive equipment and lots of experimenter time. However, human episodic memory (the most severely affected aspect of memory in AD) is different to rodent memory, which seems to be largely non-episodic. Therefore the present ways of screening for new AD treatments for AD in rodents are intrinsically unlikely to succeed. A new approach to preclinical screening would be to characterise the ADL of mice. Fortuitously, several such assays have recently been developed at Oxford, and here the three most sensitive and well-characterised are presented.Burrowing was first developed in Oxford13. It evolved from a need to develop a mouse hoarding paradigm. Most published rodent hoarding paradigms required a distant food source to be linked to the home cage by a connecting passage. This would involve modifying the home cage as well as making a mouse-proof connecting passage and food source. So it was considered whether it would be possible to put the food source inside the cage. It was found that if a container was placed on the floor it was emptied by the next morning., The food pellets were, however, simply deposited in a heap at the container entrance, rather than placed in a discrete place away from the container, as might be expected if the mice were truly hoarding them. Close inspection showed that the mice were performing digging ("burrowing") movements, not carrying the pellets in their mouths to a selected place as they would if truly hoarding them.6 Food pellets are not an essential substrate for burrowing; mice will empty tubes filled with sand, gravel, even soiled bedding from their own cage. Moreover, they will empty a full tube even if an empty one is placed next to it8. Several nesting protocols exist in the literature. The present Oxford one simplifies the procedure and has a well-defined scoring system for nest quality5. A hoarding paradigm was later developed in which the mice, rather than hoarding back to the real home cage, were adapted to living in the "home base" of a hoarding apparatus. This home base was connected to a tube made of wire mesh, the distal end of which contained the food source. This arrangement proved to yield good hoarding behaviour, as long as the mice were adapted to living in the "home base" during the day and only allowed to enter the hoarding tube at night.

The influence of travel distance on sugar loading decisions and water balance in the central place foraging ant Solenopsis invicta

One reason given for the success of social insects is their exceptional efficiency in foraging. In this study we examined the recruitment trail dynamics of sucrose foraging red imported fire ants at three distances in order to understand the limitations of their recruitment ability. Five biologically and statistically significant differences were found among several important parameters between the furthest distance (32 m) and the two closer distances (8 and 16 m): 1) Recruits leaving the food at the greatest distance had significantly more load than recruits leaving the food at the shorter distances did. 2) At the greatest distance, recruits approaching the food already had significant loads. 3)Way stations and counter-current food passing were only found when food was placed at the furthest distance. 4) Ants leaving the colony for the most distant food source were significantly smaller than recruits leaving for the two closer distances. 5) The majority of recruits returning to the colony from the furthest distance had no detectable sugars, and were also dehydrated. Our data suggest that maintaining a minimum viable level of body water appears to be more important as a limiting factor to foraging range than energetics, at least under humidity regimes that are less than saturated.

Significance of Honeybee Recruitment Strategies Depending on Foraging Distance (Hymenoptera: Apidae: Apis mellifera)

The importance of the spatial information which is communicated in the Carnolian Race of the Western Honeybee, Apis mellifera carnica (Pollmann 1879) waggle dance relative to other cues used by bees in finding food sources was investigated. The efficiency of recruitment with and without transmission of direction information in the waggle dance was quantified using artificial, plentiful unscented food sources and hives which were turned to a horizontal position to disrupt orientation of dancing bees and thereby eliminate the spatial information from dances. Transmission of location information seems to improve recruitment effect particularly at large distances. Recruitment declines more rapidly with distance if dances are disoriented, and for large distances it takes a few hours before a foraging group is established. However, this shows that even without dance information, foragers manage to recruit some bees to their food source. This process, however, is so slow that by the time a group of recruits has reached the food source, it may not be worth exploiting any more. Transmission of spatial information thus is especially important if distant food sources which often change in nectar availability are exploited.

Comparative analysis of food‐finding behavior of an herbivorous and a carnivorous land snail

Abstract. Although the olfactory capabilities of land snail tentacles have been tested by lesion studies and unilateral exposure of tentacles to specific odors, studies of a carnivorous species suggest that the anatomical similarities of herbivorous and carnivorous land snails may belie a fundamental difference in the way these structures are used to find food. Therefore, we challenged the herbivore, Anguispira alternata, and the carnivore, Haplotrema concavum, to find a stationary food source (carrot and caged young prey snail, respectively) under identical still air conditions. The herbivore traveled a significantly shorter distance to the food, even negotiating a barrier placed halfway between the snail and its food. The carnivore, on the other hand, followed a circuitous, apparently random, path to the food. Subsequent tests revealed that H. concavum readily follows prey slime trails while A. alternata seldom follows conspecific slime trails when a distant food source is available. These results are consistent with what might be expected as adaptations to the usually mobile nature of carnivore prey and the stationary nature of herbivore food plants. The ability of A. alternata to exhibit typical detour behavior is noted.

Lateral optic flow does not influence distance estimation in the desert ant Cataglyphis fortis.

The present account answers the question of whether desert ants (Cataglyphis fortis) gauge the distance they have travelled by using self-induced lateral optic-flow parameters, as has been described for bees. The ants were trained to run to a distant food source within a channel whose walls were covered with black-and-white gratings. From the food source, they were transferred to test channels of double or half the training width, and the distance they travelled before searching for home and their walking speeds were recorded. Since the animals experience different motion parallax cues when walking in the broader or narrower channels, the optic-flow hypothesis predicted that the ants would walk faster and further in the broader channels, but more slowly and less far in the narrower channels. In contrast to this expectation, neither the walking speeds nor the searching distances depended on the width or height of the channels or on the pattern wavelengths. Even when ventral-field visual cues were excluded by covering the eyes with light-tight paint, the ants were not influenced by lateral optic flow-field cues. Hence, walking desert ants do not depend on self-induced visual flow-field cues in gauging the distance they have travelled, as do flying honeybees, but can measure locomotor distance exclusively by idiothetic means.

Orientation of Aplysia californien to distant food sources

The behavior of the marine mollusc Aplysia was examined under different experimental conditions designed to determine the food searching strategy of the animals. In a small, open field tank with still water, the animals took an average of 42 min to find a piece of seaweed, even though the stimulus was never located more than 30 cm away from the animal. Observations of the animals indicated that their search was not directed, without a clear tendency towards the food, and during the course of a search, they often crawled through most of the area of the tank. The search time, the distance travelled, and the strategy of the search of the animals was similar for different types of seaweed. If animals were aroused into activity by the presence of seaweed extract, the time for them to contact a piece of odorless glass fiber paper in the open field was not significantly different than that for a piece of seaweed. The probability at which the animals contacted the seaweed, as a function of the distance travelled, resembled the detection probability determined according to a theory of random search. We thus propose that the aroused animals move in a random pattern until they are very close to the food. This strategy can be advantageous in still water since chemicals do not provide distinct gradients that can serve as cues for chemotactic orientation from distances greater than a few centimeters from the source. In a Y-maze in still water, Aplysia did not perform above chance in selecting the arm that contained the seaweed.(ABSTRACT TRUNCATED AT 250 WORDS)

GASTROPOD CHEMORECEPTION

Summary(I). Gastropods use chemoreception for a wide variety of behaviours including feeding, homing, escape from predators and a variety of social and reproductive behaviours. Chemoreception is used to locate distant food sources, and to discriminate between potential foods. Responses to chemical food stimuli result from a combination of innate and experiential factors. Gastropods use chemical cues in mucus trails to home. They also home by direct olfactory orientation. Reproductive behaviour in a variety of gastropods appears to involve chemical cues. Evidence exists for pheromones controlling aggregation and mating. Numerous gastropods use chemical cues to avoid or escape from predators.(2). Amino acids appear as likely candidates for attractants and phagostimulants for gastropod feeding. Macromolecules are probably also involved. Amino acids have also been shown to stimulate reproductive behaviours in certain gastropods, thus suggesting a pheromonal function. However, the significance of this finding to the behaviour of the organisms in the field has yet to be evaluated. Saponins have been implicated as the active substances found in sea stars that elicit escape responses of marine gastropods. Choline esters may play a homologous role in gastropod—prey and gastropod‐predator interactions.(3). Gastropods can apparently use a number of different methods to orient to olfactory cues. These include anemotaxis or rheotaxis, klinotaxis and tropotaxis.(4). The major chemosensory organs of gastropods have been identified. They include the anterior and posterior tentacles and lips of terrestrial pulmonates; the cephalic tentacles, the lips and buccal cavity lining, and possibly the osphradium of aquatic pulmonates; the cephalic and mantle tentacles, the anterior margin of the foot, the siphon tip, and the osphradium of prosobranchs; and the rhinophores, tentacles, oral veil and osphradium of opisthobranchs.(5). Many of the organs named above have been examined by both light and electron microscopy. The most common anatomical organization includes bipolar primary sensory cells with cell bodies located subepithelially, and a distal dendrite extending to the free surface. Often a peripheral ganglion is located deep to the sensory epithelium. It is unclear whether axons of the sensory cells project directly to the central ganglion or by way of interneurones located in the peripheral ganglia.(6). The dendritic specializations of the sensory cells vary considerably. Most bear cilia or a combination of cilia and microvilli. The functional significance of the variation in the types of sensory endings is unknown, although the chemosensory epithelia also respond to other sensory modalities, and it is often difficult to ascribe any one cell type to any one modality. Species‐specific variations may also complicate the picture.(7). Prospects for and importance of future studies on gastropod chemoreception are discussed.

The honey-bee “dance language” hypothesis and the foundations of biology and behavior

The ability of new bees to arrive at distant food sources by use of odors, and not “dance language” information, has long been proven. Their ability to use “dance language” information has not yet been proven. Acceptance of the “dance language” hypothesis as fact has already led to a questioning of the validity of Morgan's Canon which underlies the modern science of animal behavior. The questioning is not surprising. The final acceptance of the “dance language” hypothesis by its supporters, is based only on two recent presumed proofs by Schricker, and by Gould. An analysis of these proofs demonstrates, however, that they cast doubt not only on Morgan's Canon, but also on the theory of evolution which underlies the modern science of biology. Fortunately the claims of Schricker's proof are experimentally refuted by his own control experiment. The claims of Gould's proof are experimentally refuted by a control experiment which precedes the experiments underlying the proof. The credibility of Morgan's Canon and of the theory of evolution are, thus, fully restored.

ADAPTATION, CONSTRAINT, AND COMPROMISE IN AVIAN POSTNATAL DEVELOPMENT

Summary1. This paper discusses factors that influence the evolution of growth rate and determine its variation among species of birds. Growth rate is related to evolutionary fitness through the use of time, energy, and nutrients. In addition, balances between factors favouring rapid growth and those favouring slow growth may be investigated directly by experiment and by comparative observation.2. David Lack (1968) proposed that the growth rate of the young is the optimum balance between selection for rapid growth to reduce the vulnerable period of development and selection for slow growth to reduce the energy requirements of the young.3. To test Lack's hypothesis, the growth rates of birds, estimated by fitting sigmoid equations to curves relating weight to age, were surveyed widely from the literature. Among all species examined, growth rate was inversely related to adult weight. Among birds of similar size, most variation in growth rate was related to the degree of maturity of the neonate. Altricial chicks, which depend upon their parents for food and warmth, grow more rapidly than precocial chicks, which are self‐sufficient shortly after hatching. Lack's hypothesis, which predicts a direct relationship between growth rate and mortality rate, was not supported.4. I propose that the key to understanding variation in growth rate among birds lies in the balance between rate of cell proliferation or cell growth, on one hand, and acquisition of mature function, on the other. This idea is consistent with principles of cellular and developmental biology. It is supported by comparisons of (a) the neonates of different species, (b) the individual over the course of the developmental period, and (c) tissues whose use is acquired at different stages of development, wherein more mature individuals or tissues grow more slowly than those with less developed function.5. Species of birds that are classified as semi‐precocial develop precocially but grow rapidly. Although these seemingly violate the general rule relating growth rate to precocity, a closer inspection of their development reveals that they too support the rule. In the Common Tern, the legs, which are the key organ in precocial development, grow at the expected slow rate. The body as a whole grows rapidly because the growth increment of the legs is small and their growth is completed quickly.6. Growth rates of precocial birds do not decrease abruptly at hatching. This points more to gradual tissue differentiation than to the pattern of procurement and allocation of energy as the primary control for growth rate.7. Precocious development is favoured when the chicks are capable of self‐feeding or when food supplies are distant from the next site and travelling time between one and the other is long. Precocity of the neonates frees both parents to feed at a distant food source.8. Some species having diets with low levels of protein or other nutrients may grow slowly in order to match nutrient requirements to their availability in the diet. This pattern is indicated especially among the Procellariiformes, which feed an oily diet to their young, and also among tropical fruit‐eating birds.9. Some tropical, pelagically‐feeding sea‐birds that rear only one offspring at a time may not be able to procure food sufficient to support rapid chick growth. Alternative explanations for slow growth among these species include difficulty in obtaining essential nutrients and more precocious development of activity than in related species having more rapid growth.