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Tubulin Evolution In Insects: Gene Duplication And Subfunctionalization Provide Specialized Isoforms In A Functionally Constrained Gene Family, Mark G. Nielsen, Sudhindra R. Gadagkar, Lisa Gutzwiller
Tubulin Evolution In Insects: Gene Duplication And Subfunctionalization Provide Specialized Isoforms In A Functionally Constrained Gene Family, Mark G. Nielsen, Sudhindra R. Gadagkar, Lisa Gutzwiller
Mark G. Nielsen
Background: The completion of 19 insect genome sequencing projects spanning six insect orders provides the opportunity to investigate the evolution of important gene families, here tubulins. Tubulins are a family of eukaryotic structural genes that form microtubules, fundamental components of the cytoskeleton that mediate cell division, shape, motility, and intracellular trafficking. Previous in vivo studies in Drosophila find a stringent relationship between tubulin structure and function; small, biochemically similar changes in the major alpha 1 or testis-specific beta 2 tubulin protein render each unable to generate a motile spermtail axoneme. This has evolutionary implications, not a single non-synonymous substitution is …
The Global Decline Of Reptiles, Deja’ Vu Amphibians, J. Whitfield Gibbons, David E. Scott, Travis J. Ryan, Kurt A. Buhlmann, Tracey D. Tiuberville, Brian S. Metts, Judith L. Greene, Tony Mills, Yale Leiden, Sean Poppy, Christopher T. Winne
The Global Decline Of Reptiles, Deja’ Vu Amphibians, J. Whitfield Gibbons, David E. Scott, Travis J. Ryan, Kurt A. Buhlmann, Tracey D. Tiuberville, Brian S. Metts, Judith L. Greene, Tony Mills, Yale Leiden, Sean Poppy, Christopher T. Winne
Travis J. Ryan
Reptile species are declining on a global scale. Six significant threats to reptile populations are habitat loss and degradation, introduced invasive species, environmental pollution, disease, unsustainable use, and global climate change.
Effects Of Hydroperiod On Metamorphosis In Rana Sphenocephala, Travis J. Ryan, Christopher T. Winne
Effects Of Hydroperiod On Metamorphosis In Rana Sphenocephala, Travis J. Ryan, Christopher T. Winne
Travis J. Ryan
Hydroperiod, the time a temporary pond holds water, is an important factor influencing recruitment in amphibian populations and structuring amphibian communities. We conducted an experiment to test the effect of hydroperiod on metamorphic traits of the southern leopard frog (Rana sphenocephala), a common amphibian in the southeastern United States. We reared larval R. sphenocephala in artificial ponds at a density of 32 larvae per tank (initial volume = approximately 650 liter). We dried the tanks according to natural patterns, using three different hydroperiods (60, 75 and 90 d). Experimental hydroperiods had a significant effect on the number of metamorphs and …
Hatching Asynchrony, Survival, And The Fitness Of Alternative Adult Morphs In Ambystoma Talpoideum, Travis J. Ryan
Hatching Asynchrony, Survival, And The Fitness Of Alternative Adult Morphs In Ambystoma Talpoideum, Travis J. Ryan
Travis J. Ryan
The mole salamander, Ambystoma talpoideum, exhibits both aquatic (gilled) and terrestrial (metamorphosed) adult morphologies. Previous studies have shown the existence of body-size advantages associated with the terrestrial morph in A. talpoideum and other polymorphic salamanders (e.g., A. tigrinum). However, aquatic adult A. talpoideum mature at a younger age and often breed earlier than terrestrial adults. We tested the hypothesis that early maturation and reproduction in aquatic adults increase fitness (irrespective of body size). We reared larval A. talpoideum in mesocosms and varied the timing of hatching, with early-hatching larvae representing the offspring from early-breeding aquatic adults, and late-hatching larvae representing …
Monitoring Herpetofauna In A Managed Forest Landscape: Effects Of Habitat Types And Census Techniques, Travis J. Ryan, Thomas Philippi, Yale A. Leiden, Michael E. Dorcas, T. Bently Wigley, J. Whitfield Gibbons
Monitoring Herpetofauna In A Managed Forest Landscape: Effects Of Habitat Types And Census Techniques, Travis J. Ryan, Thomas Philippi, Yale A. Leiden, Michael E. Dorcas, T. Bently Wigley, J. Whitfield Gibbons
Travis J. Ryan
We surveyed the herpetofaunal (amphibian and reptile) communities inhabiting five types of habitat on a managed landscape. We conducted monthly surveys during 1997 in four replicate plots of each habitat type using several different methods of collection. Communities of the two wetland habitats (bottomland wetlands and isolated upland wetlands) were clearly dissimilar from the three terrestrial communities (recent clearcut, pine plantation, and mixed pine–hardwood forest). Among the three terrestrial habitats, the total herpetofaunal communities were dissimilar (P<0.10), although neither faunal constituent group alone (amphibians and squamate reptiles) varied significantly with regard to habitat. Three survey techniques used in the …
Sub-Lethal Effects Of 2,4-D Exposure On Golf Course Amphibians, Travis J. Ryan, Catherine M. Scott, Brooke A. Douthitt
Sub-Lethal Effects Of 2,4-D Exposure On Golf Course Amphibians, Travis J. Ryan, Catherine M. Scott, Brooke A. Douthitt
Travis J. Ryan
Amphibians are among the most common vertebrates inhabiting golf courses. They are frequent inhabitants of golf course wetlands, where they are likely to be episodically exposed to small quantities of herbicides in proper golf course maintenance. The goal of this study was to investigate whether amphibian larvae subjected to sublethal concentrations of a common herbicide used in golf course maintenance would likely result in significant life history, locomotor, or behavioral effects (e.g., changes in growth, swimming speed, or feeding ability). For these experiments we selected as models tadpoles of the southern leopard frog (Rana sphenocephala) and the herbicide 2,4- dichlorophenoxyacetic …
Intraspecific Heterochrony And Life History Evolution: Decoupling Somatic And Sexual Development In A Facultatively Paedomorphic Salamander, Travis J. Ryan, Raymond D. Semlitsch
Intraspecific Heterochrony And Life History Evolution: Decoupling Somatic And Sexual Development In A Facultatively Paedomorphic Salamander, Travis J. Ryan, Raymond D. Semlitsch
Travis J. Ryan
Morphological features such as size and shape are the most common focus in studies of heterochronic change. Frequently, these easily observed and measured features are treated as a major target of selection, potentially ignoring traits more closely related to fitness. We question the primacy of morphological data in studies of heterochrony, and instead suggest that principal sources of fitness, such as life history characteristics, are not only the chief targets of selection, but changes in them may necessitate changes in other (subordinate) elements of the organism. We use an experimental approach to investigate the timing of metamorphosis and maturation in …
Trials Of The Urban Ecologist, Rebecca W. Dolan, Tim Carter, Travis J. Ryan, Carmen M. Salsbury, Thomas E. Dolan, Marjorie Hennessy
Trials Of The Urban Ecologist, Rebecca W. Dolan, Tim Carter, Travis J. Ryan, Carmen M. Salsbury, Thomas E. Dolan, Marjorie Hennessy
Travis J. Ryan
A group of scientists describe some of the obstacles encountered and insights gained while carrying out ecological research in and around the city of Indianapolis.
The Molecular Dimension Of Microbial Species: 3. Comparative Genomics Of Synechococcus Strains With Different Light Responses And In Situ Diel Transcription Patterns Of Associated Putative Ecotypes In The Mushroom Spring Microbial Mat, Millie Olsen, Shane Nowack, Jason Wood, Eric Becraft, Kurt Labutti, Anna Lipzen, Joel Martin, Wendy Shackwitz, Douglas Rusch, Frederick Cohan, Donald Bryant, David Ward
The Molecular Dimension Of Microbial Species: 3. Comparative Genomics Of Synechococcus Strains With Different Light Responses And In Situ Diel Transcription Patterns Of Associated Putative Ecotypes In The Mushroom Spring Microbial Mat, Millie Olsen, Shane Nowack, Jason Wood, Eric Becraft, Kurt Labutti, Anna Lipzen, Joel Martin, Wendy Shackwitz, Douglas Rusch, Frederick Cohan, Donald Bryant, David Ward
Frederick M. Cohan
Genomes were obtained for three closely related strains of Synechococcus that are representative of putative ecotypes that predominate at different depths in the 1 mm-thick, upper-green layer in the 60°C mat of Mushroom Spring, Yellowstone National Park, and exhibit different light adaptation and acclimation responses. The genomes were compared to the published genome of a previously obtained, closely related strain from a neighboring spring, and differences in both gene content and orthologous gene alleles between high-light-adapted and low-light-adapted strains were identified. Evidence of genetic differences that relate to adaptation to light intensity and/or quality, CO2 uptake, nitrogen metabolism, organic carbon …
Exploring Behavioral And Hormonal Flexibility Across Light Environments In Guppies From Low-Predation Populations, Gita R. Kolluru
Exploring Behavioral And Hormonal Flexibility Across Light Environments In Guppies From Low-Predation Populations, Gita R. Kolluru
Gita R. Kolluru
No abstract provided.
The Molecular Dimension Of Microbial Species: 1. Ecological Distinctions Among, And Homogeneity Within, Putative Ecotypes Of Synechococcus Inhabiting The Cyanobacterial Mat Of Mushroomspring,Yellowstone National Park, Eric D. Becraft, Jason M. Wood, Douglas B. Rusch, Michael Kuhl, Sheila Jensen, Donald A. Bryant, David W. Roberts, Frederick M. Cohan, David M. Ward
The Molecular Dimension Of Microbial Species: 1. Ecological Distinctions Among, And Homogeneity Within, Putative Ecotypes Of Synechococcus Inhabiting The Cyanobacterial Mat Of Mushroomspring,Yellowstone National Park, Eric D. Becraft, Jason M. Wood, Douglas B. Rusch, Michael Kuhl, Sheila Jensen, Donald A. Bryant, David W. Roberts, Frederick M. Cohan, David M. Ward
Frederick M. Cohan
Based on the Stable Ecotype Model, evolution leads to the divergence of ecologically distinct populations (e.g., with different niches and/or behaviors) of ecologically interchangeable membership. In this study, pyrosequencing was used to provide deep sequence coverage of Synechococcus psaA genes (encoding a photosystem I reaction center protein subunit) and transcripts over a large number of habitat types in the Mushroom Spring microbial mat. Putative ecological species (putative ecotypes), which were predicted by an evolutionary simulation based on the Stable Ecotype Model (Ecotype Simulation), exhibited distinct distributions relative to temperature-defined positions in the effluent channel and vertical position in the upper …
Calf Development: Most Births At Night, Robert H.I. Dale
Calf Development: Most Births At Night, Robert H.I. Dale
Robert H. I. Dale
For many years, field researchers studying both African (Loxodonta africana) and Asian (Elephas maximas) elephants have indicated that they have observed relatively few births in situ, suggesting that most elephant dams give birth at night. For example, according to Cynthia Moss, "Possibly the majority of births occur at night and perhaps those that do take place in the daytime happen in secluded places" (1988, p. 151). Others, for example, Clive Spinage, have referred to "the old beliefs that the cows retreated to 'calving grounds' or that birth took place at night." (Spinage, 1994, p. 90). Although observers in several areas …
Tribute: In Memoriam, Jeheskel "Hezy" Shoshani, Robert H.I. Dale
Tribute: In Memoriam, Jeheskel "Hezy" Shoshani, Robert H.I. Dale
Robert H. I. Dale
The elephant community has lost one of its great ambassadors, Dr. Jeheskel "Hezy" Shoshani, at the age of 65. A wave of condolences and testimonials from colleagues and friends around the world followed his death on May 21, 2008.
History: The Birth Of "America" In 1882, Robert H.I. Dale
History: The Birth Of "America" In 1882, Robert H.I. Dale
Robert H. I. Dale
This article concerns a New York Times story about the birth of the female Asian elephant calf, named America, at the winter headquarters of the "Greatest Show on Earth" in Bridgeport, Connecticut on February 2, 1882. Phineas T. Barnum, one of the owners of the show, and one prone to self-aggrandizing bluster, claimed that America was the second elephant ever born in captivity. America was born only to months before the arrival in New York of the most famous circus elephant of all time, Jumbo, on Easter Sunday, 1882, and only two years before the origin of a small wagon …
Variations In Radial Maze Performance Under Different Levels Of Food And Water Deprivation, Robert H.I. Dale, William A. Roberts
Variations In Radial Maze Performance Under Different Levels Of Food And Water Deprivation, Robert H.I. Dale, William A. Roberts
Robert H. I. Dale
Four groups of rats were tested on an eight-arm radial maze under a free-choice procedure. The subjects were maintained at either 80% or 100% of their preexperimental free-feeding weights through restricted access to either food or water. Water-deprived subjects received water in the maze; food-deprived subjects received food. Water-deprived subjects learned the task faster than food-deprived subjects. The four groups developed different response patterns. These were measured by the mean transition size, the average angular distance (in 45° units) between consecutively chosen arms. Rats foraging for food and water developed different search strategies, with water-deprived subjects exhibiting lower mean transition …
Similarities Between Human And Animal Spatial Memory: Item And Order Information, Robert H.I. Dale
Similarities Between Human And Animal Spatial Memory: Item And Order Information, Robert H.I. Dale
Robert H. I. Dale
Human subjects, sitting at the center of a circle of eight lights, were tested on analogues of radial-maze item-recognition (Roberts & Smythe, 1979) and order-recognition (Kesner & Novak, 1982) tasks. Subjects in the item-recognition condition saw a list of seven lights, and then the nonlist (eighth) light was tested against the first, fourth, or seventh light from the list. The sub- jects were required to point toward the non list light. Subjects in the order-recognition condition saw a series of eight lights, followed by a test of the first and second, fourth and fifth, or seventh and eighth serial positions. …
Spatial And Temporal Response Patterns On The Eight-Arm Radial Maze, Robert H.I. Dale
Spatial And Temporal Response Patterns On The Eight-Arm Radial Maze, Robert H.I. Dale
Robert H. I. Dale
Six maze-experienced hooded rats were timed during five trials on which they collected water from all arms of an eight-arm radial maze, then made five more choices. All subjects frequently exhibited a “task-completion pause:” The subjects rarely spent more than 1 sec in the center of the maze between choices until they had entered all eight arms, then stopped in the center of the maze. In contrast, the time spent in each arm gradually increased until all of the water had been obtained, then decreased slightly. Four subjects began every trial by choosing eight consecutive adjacent arms. The task-completion pause …
Some Children's Books About Elephants (1987-1997), Robert H.I. Dale
Some Children's Books About Elephants (1987-1997), Robert H.I. Dale
Robert H. I. Dale
Literature review of children's books about elephants from the Indianapolis Public Library.
Spatial Memory In Pigeons On A Four-Arm Radial Maze, Robert H.I. Dale
Spatial Memory In Pigeons On A Four-Arm Radial Maze, Robert H.I. Dale
Robert H. I. Dale
Pigeon spatial memory was examined using a four-arm radial maze. The maze had four arms, spaced at 90° intervals, extending radially from a central choice area. Subjects were forced into three arms, then permitted two choices to enter the remaining ann. Five subjects chose accurately (90% correct) with delays of 5 min or less, their choices depended on extramaze cues, and the food in the target arm provided no essential cues. After an incorrect first choice, subjects' second choices were more accurate than chance. These data suggest that, while spatial memory has many similar characteristics in rats and pigeons, pigeon …
The Locomotor Kinematics Of Asian And African Elephants: Changes With Speed And Size, John R. Hutchinson, Delf Schwerda, Daniel J. Famini, Robert H.I. Dale, Martin S. Fischer, Rodger Kram
The Locomotor Kinematics Of Asian And African Elephants: Changes With Speed And Size, John R. Hutchinson, Delf Schwerda, Daniel J. Famini, Robert H.I. Dale, Martin S. Fischer, Rodger Kram
Robert H. I. Dale
For centuries, elephant locomotion has been a contentious and confusing challenge for locomotion scientists to understand, not only because of technical difficulties but also because elephant locomotion is in some ways atypical of more familiar quadrupedal gaits. We analyzed the locomotor kinematics of over 2400 strides from 14 African and 48 Asian elephant individuals (body mass 116-4632 kg) freely moving over ground at a 17-fold range of speeds, from slow walking at 0.40 m s-1 to the fastest reliably recorded speed for elephants, 6.8 m s-1. These data reveal that African and Asian elephants have some subtle differences in how …
A Survey Of The Management And Development Of Captive African Elephant (Loxodonta Africana) Calves: Birth To Three Months Of Age, Nicole L. Kowalski, Robert H.I. Dale, Christa L. H. Mazur
A Survey Of The Management And Development Of Captive African Elephant (Loxodonta Africana) Calves: Birth To Three Months Of Age, Nicole L. Kowalski, Robert H.I. Dale, Christa L. H. Mazur
Robert H. I. Dale
We used four surveys to collect information about the birth, physical growth, and behavioral development of 12 African elephant calves born in captivity. The management of the birth process and neonatal care involved a variety of standard procedures. All of the calves were born at night, between 7PM and 7AM. The calves showed a systematic progression in behavioral and physical development, attaining developmental milestones at least a quickly as calves in situ. This study emphasized birth-related events, changes in the ways that calves used their trunks, first instances of behaviors, and interactions of the calves with other, usually adult, elephants. …
The Relative Attenuation Of Self-Stimulation, Eating And Drinking Produced By Dopamine-Receptor Blockade, E. T. Rolls, B. J. Rolls, P. H. Kelly, S. G. Shaw, R. J. Wood, Robert H.I. Dale
The Relative Attenuation Of Self-Stimulation, Eating And Drinking Produced By Dopamine-Receptor Blockade, E. T. Rolls, B. J. Rolls, P. H. Kelly, S. G. Shaw, R. J. Wood, Robert H.I. Dale
Robert H. I. Dale
Spiroperidol, which blocks dopamine (DA) receptors, attenuated self-stimulation of the nucleus accumbens, septal area, hippocampus, anterior hypothalamus and ventral tegmental area. Dopamine is thus involved in self-stimulation of many sites (in addition to the lateral hypothalamus). The attenuation was not a simple motor impairment of the speed of bar-pressing in that the nucleus accumbens and septal self-stimulation rates were lower than those in treated animals self-stimulating at other sites (Experiment 1). Feeding was partly attenuated, and drinking was much less attenuated by the spiroperidol. Since the rats bar-pressed for brain- stimulation reward, chewed pellets to eat, and licked a tube …
Limitations On Spatial Memory In Mice, Robert H.I. Dale, Martin Bedard
Limitations On Spatial Memory In Mice, Robert H.I. Dale, Martin Bedard
Robert H. I. Dale
Rats have an impressive ability to remember locations they have visited. Two experiments used an eight-arm radial maze to determine whether mice showed two important characteristics of this spatial memory: its durability, and its dependence on stimuli outside the maze (extreme stimuli). In Experiment 1, food-deprived mice were allowed to eat from four of the eight arms of the maze then, after delays of 5 sec, 1 min, or 5 min, they were permitted to choose the remaining arms. Choice accuracy declined significantly with the longer delays, but always remained above chance. In Experiment 2, the maze was rotated 180° …
The Hippocampus As Episodic Encoder: Does It Play Tag?, Robert H.I. Dale
The Hippocampus As Episodic Encoder: Does It Play Tag?, Robert H.I. Dale
Robert H. I. Dale
Rawlins’s characterization of the hippocampus as a “high-capacity, immediate-term memory store” captures the essential idea in a number of previous models. For example, Gaffan (1974), Gray (1984), Hirsh (1980), Kesner (Bierley, Kesner & Novak 1983), Olton (Olton, Becker & Handelmann 1979), Solomon (1980), and Winocur (1980) all agree that hippocampal animals show memory deficits when required to identify, for whatever reason, one specific event out of a list of recent events. Although these authors disagree on a number of details, Rawlins has identified their models common ground, the core of each model. (It is only fair to note that Gaffan …
Remembrance Of Places Lasts: Proactive Inhibition And Patterns Of Choice In Rat Spatial Memory, William A. Roberts, Robert H.I. Dale
Remembrance Of Places Lasts: Proactive Inhibition And Patterns Of Choice In Rat Spatial Memory, William A. Roberts, Robert H.I. Dale
Robert H. I. Dale
A series of experiments was carried out to evaluate the notion that rats given a sequence of massed daily trials on the radial maze reset working memory at the end of each trial by deleting its contents. Although curves presented by D. S. Olton [Scientific American, 1977, 236, 82-98: In S. H. Hulse, H. Fowler, & W. K. Honig (Eds.), Cognitive processes in animal behavior, 1978, Hillsdale, N.J.: Erlbaum] show that rats return to errorless performance at the beginning of each trial after the first, the fact that accuracy falls less rapidly over choices on Trial 1 …
Radial-Maze Performance In The Rat Following Lesions Of Posterior Neocortex, Melvyn A. Goodale, Robert H.I. Dale
Radial-Maze Performance In The Rat Following Lesions Of Posterior Neocortex, Melvyn A. Goodale, Robert H.I. Dale
Robert H. I. Dale
The present experiment was designed to investigate the role of posterior neocortex (areas 17, 18 and 18a) in the maintenance of performance on the radial maze. Following training to criterion on the 8-arm radial maze, rats received either sham operations, bilateral eye enucleations, lesions of posterior neocortex, or combined enucleations and lesions of posterior neocortex. While the enucleated animals with intact brains showed a slight, but significant performance decrement relative to the sham-operated group, the other two groups, with lesions of areas 17, 18 and 18a, each showed a massive deficit. This large deficit was observed even in the group …
Parallel-Arm Maze Performance Of Sighted And Blind Rats: Spatial Memory And Maze Structure, Robert H.I. Dale
Parallel-Arm Maze Performance Of Sighted And Blind Rats: Spatial Memory And Maze Structure, Robert H.I. Dale
Robert H. I. Dale
Sighted and peripherally blinded groups of rats learned to obtain a small reward from each arm of an eight-arm parallel maze, and a sighted group was similarly trained on a radial maze. The parallel-sighted and parallel-blind groups were equally slow, and much slower than the radial-sighted group, to attain criterion performance. The three groups shared several response characteristics: selectively avoiding the most recently entered arms, frequently choosing adjacent arms, and an absence of 'spatial generalization' among the arms. The findings support a simple model proposing how subjects identify and choose among the maze-arms.
Dynamic Effects Of Food Magnitude On Interim-Terminal Interaction, Alliston K. Reid, Robert H.I. Dale
Dynamic Effects Of Food Magnitude On Interim-Terminal Interaction, Alliston K. Reid, Robert H.I. Dale
Robert H. I. Dale
We tested the assumption of a facilitatory relation between periodic food presentation and schedule-induced drinking by examination of (a) elicited drinking, (b) drinking in anticipation of food delivery, and (c) possible indirect effects of food delivery on drinking. We exposed rats to a fixed-time 60-second schedule in which interfood intervals ended in either one or four food pellets with equal probability. In Phases 1 and 3, a stimulus signaled the magnitude of upcoming food presentation. In Phase 2, the stimulus was eliminated. Changes in drinking and "head-in-feeder" distributions within interfood intervals demonstrated that head-in-feeder was controlled directly by food presentation, …
Interactions Between Response Stereotypy And Memory Strategies On The Eight-Arm Radial Maze, Robert H.I. Dale, Nancy K. Innis
Interactions Between Response Stereotypy And Memory Strategies On The Eight-Arm Radial Maze, Robert H.I. Dale, Nancy K. Innis
Robert H. I. Dale
Three groups of water-deprived rats collected water from the ends of the 8 arms of an 8-arm radial maze. Sighted subjects, and subjects blinded either with or without pre-enucleation experience on the radial maze, all retrieved the water efficiently. Most of the subjects exhibited the same response stereotypy, regularly choosing 8 adjacent arms of the maze, then stopping in the center of the maze. The strategies underlying this performance were analyzed by interrupting trials and rotating the maze 180° after the subject had made 3 choices. Sighted subjects depended on extramaze stimuli, naive-blind subjects depended on intramaze stimuli and experienced-blind …
Concurrent Drinking By Pigeons On Fixed-Interval Reinforcement Schedules, Robert H.I. Dale
Concurrent Drinking By Pigeons On Fixed-Interval Reinforcement Schedules, Robert H.I. Dale
Robert H. I. Dale
Three experienced pigeons were exposed to at least ten consecutive 100-min sessions on each of three food-reinforced fixed-interval (FI) schedules: FI 50-sec, FI 100-sec and FI 200-sec. Water was freely available. Drinking was largely confined to the first third of each fixed interval, and the mean sessional water intake was directly related to the food-reinforcement rate for each animal. The animals drank very quickly, i.e., 3-4 ml/sec, but the drinking bouts were brief, i.e., 0.8-1.4 sec, and infrequent, i.e., 2-5/hr. The parameters describing concurrent drinking in the pigeon are strikingly different from those describing rats’ drinking under similar reinforcement schedules, …