Douglas Elliffe

Complex cognition and behavioural innovation in New Caledonian crows

Apes, corvids and parrots all show high rates of behavioural innovation in the wild. However, it is unclear whether this innovative behaviour is underpinned by cognition more complex than simple learning mechanisms. To investigate this question we presented New Caledonian crows with a novel three-stage metatool problem. The task involved three distinct stages: (i) obtaining a short stick by pulling up a string, (ii) using the short stick as a metatool to extract a long stick from a toolbox, and finally (iii) using the long stick to extract food from a hole. Crows with previous experience of the behaviours in stages 1–3 linked them into a novel sequence to solve the problem on the first trial. Crows with experience of only using string and tools to access food also successfully solved the problem. This innovative use of established behaviours in novel contexts was not based on resurgence, chaining and conditional reinforcement. Instead, the performance was consistent with the transfer of an abstract, causal rule: ‘out-of-reach objects can be accessed using a tool’. This suggests that high innovation rates in the wild may reflect complex cognitive abilities that supplement basic learning mechanisms.

Correlation of cellular changes and spatial memory during aging in rats.

We describe neuronal density, neuroplasticity and vascular remodelling and their association with spatial memory in young (4-6 months), middle-aged (9-11 months) and aged (18-20 months) rats of both genders. The neuronal density was reduced in the hippocampus of middle-aged and aged rats, particularly in male rats. However the loss of spatial memory investigated using the Morris water maze, T-maze and 8-radial arm maze tests was found only in the aged groups. The data suggested a pre-symptomatic period of pathological brain aging. Surprisingly, the middle-aged groups showed an elevation of glutamate-decarboxylase immunoreactive neurons in the hippocampus and the striatum, an increase of dopamine output in the striatum and enhanced vascular remodelling in the hippocampus when compared with the young and, in some cases, aged groups. Together, the data suggest that the loss of neurons during midlife may stimulate and enhance neuronal plasticity and vascular remodelling. These compensatory responses to initial neuronal degeneration may play a role in delaying impending memory loss during the pre-symptomatic period of pathological brain aging.

On the joint control of preference by time and reinforcer-ratio variation

Five pigeons were trained in a procedure in which, with a specified probability, food was either available on a fixed-interval schedule on the left key, or on a variable-interval schedule on the right key. In Phase 1, we arranged, with a probability of 0.5, either a left-key fixed-interval schedule or a right-key variable-interval 30s, and varied the value of the fixed-interval schedule from 5s to 50s across 5 conditions. In Phase 2, we arranged either a left-key fixed-interval 20-s schedule or a right-key variable-interval 30-s schedule, and varied the probability of the fixed-interval schedule from 0.05 to 1.0 across 8 conditions. Phase 3 always arranged a fixed-interval schedule on the left key, and its value was varied over the same range as in Phase 1. In Phase 1, overall preference was generally toward the variable-interval schedule, preference following reinforcers was initially toward the variable-interval schedule, and maximum preference for the fixed-interval schedule generally occurred close to the arranged fixed-interval time, becoming relatively constant thereafter. In Phase 2, overall left-key preference followed the probability of the fixed-interval schedule, and maximum fixed-interval choice again occurred close to the fixed-interval time, except when the fixed-interval probability was 0.1 or less. The pattern of choice following reinforcers was similar to that in Phase 1, but the peak fixed-interval choice became more peaked with higher probabilities of the fixed interval. Phase 3 produced typical fixed-interval schedule responding. The results are discussed in terms of reinforcement effects, timing in the context of alternative reinforcers, and generalized matching. These results can be described by a quantitative model in which reinforcer rates obtained at times since the last reinforcer are distributed across time according to a Gaussian distribution with constant coefficient of variation before the fixed-interval schedule time, changing to extended choice controlled by extended reinforcer ratios beyond the fixed-interval time. The same model provides a good description of response rates on single fixed-interval schedules.

EMERGENT STIMULUS RELATIONS DEPEND ON STIMULUS CORRELATION AND NOT ON REINFORCEMENT CONTINGENCIES

We aimed to investigate whether novel stimulus relations would emerge from stimulus correlations when those relations explicitly conflicted with reinforced relations. In a symbolic matching-to-sample task using kanji characters as stimuli, we arranged class-specific incorrect comparison stimuli in each of three classes. After presenting either Ax or Cx stimuli as samples, choices of Bx were reinforced and choices of Gx or Hx were not. Tests for symmetry, and combined symmetry and transitivity, showed the emergence of three 3-member (AxBxCx) stimulus classes in 5 of 5 human participants. Subsequent tests for all possible emergent relations between Ax, Bx, Cx and the class-specific incorrect comparisons Gx and Hx showed that these relations emerged for 4 of 5 the participants after extended overtraining of the baseline relations. These emergent relations must have been based on stimulus-stimulus correlations, and were not properties of the trained discriminated operants, because they required control by relations explicitly extinguished during training. This result supports theoretical accounts of emergent relations that emphasize stimulus correlation over operant contingencies.

A model for food and stimulus changes that signal time-based contingency changes.

When the availability of reinforcers depends on time since an event, time functions as a discriminative stimulus. Behavioral control by elapsed time is generally weak, but may be enhanced by added stimuli that act as additional time markers. The present paper assessed the effect of brief and continuous added stimuli on control by time-based changes in the reinforcer differential, using a procedure in which the local reinforcer ratio reversed at a fixed time after the most recent reinforcer delivery. Local choice was enhanced by the presentation of the brief stimuli, even when the stimulus change signalled only elapsed time, but not the local reinforcer ratio. The effect of the brief stimulus presentations on choice decreased as a function of time since the most recent stimulus change. We compared the ability of several versions of a model of local choice to describe these data. The data were best described by a model which assumed that error in discriminating the local reinforcer ratio arose from imprecise discrimination of reinforcers in both time and space, suggesting that timing behavior is controlled not only by discrimination elapsed time, but by discrimination of the reinforcer differential in time.

DIVIDED STIMULUS CONTROL: A REPLICATION AND A QUANTITATIVE MODEL

Four pigeons were trained on a conditional discrimination. The conditional stimuli were compounds of pairs of stimuli from two different dimensions, fast versus slow cycles of red or green stimuli, and short- versus long-duration presentations of these cycles. Across conditions, the probability of reinforcers for correctly responding to each dimension was varied from 0 to 1. Discriminability, measured by log d, for stimuli on a dimension increased as the relative frequency of reinforcers for that dimension increased, replicating the results of Shahan and Podlesnik (2006). Two further conditions showed that discriminability between stimuli on each dimension was unaffected by whether the stimuli on the other dimension varied or were constant. Finally, maximal discriminability was unchanged in a redundant-relevant cues condition in which either of the stimuli comprising a compound signaled the same correct response. Davison and Nevins (1999) model provided an excellent quantitative account of the effect of relative reinforcer frequency on discriminability, and thus of the way in which divided stimulus control is itself controlled by relative reinforcement.

Four-alternative choice violates the constant-ratio rule

Six pigeons responded on a four-key concurrent variable-interval schedule in which a 27:9:3:1 distribution of reinforcers between the keys changed every 10 reinforcers. Their behaviour quickly came under the control of this changing four-way reinforcer ratio. However, preference between a pair of keys depended not only on the relative reinforcer rates on those keys, but also on the absolute levels of those rates. This contradicts the constant-ratio rule that underpins the matching approach to choice, but is predicted by a contingency-discriminability model that assumes that organisms may occasionally misattribute reinforcers to a response that did not produce them. Reinforcers produced strong preference pulses, or transient increases in responding on the just-reinforced key. Despite accurate tracking of the reinforcer ratio, reinforcers obtained late in components and from leaner keys still produced strong pulses, suggesting both extended and local control of behaviour. Patterns of switching between keys were graded and similarly controlled by the reinforcer rates on each key. Whether considered in terms of switching, local preference pulses, or extended preference, behaviour was controlled by a rapidly changing four-way reinforcer ratio in a graduated, continuous manner that is unlikely to be explained by a simple heuristic such as fix-and-sample.

Stimulus–reinforcer relations established during training determine resistance to extinction and relapse via reinstatement

The baseline rate of a reinforced target response decreases with the availability of response-independent sources of alternative reinforcement; however, resistance to disruption and relapse increases. Because many behavioral treatments for problem behavior include response-dependent reinforcement of alternative behavior, the present study assessed whether response-dependent alternative reinforcement also decreases baseline response rates but increases resistance to extinction and relapse. We reinforced target responding at equal rates across two components of a multiple schedule with pigeons. We compared resistance to extinction and relapse via reinstatement of (1) a target response trained concurrently with a reinforced alternative response in one component with (2) a target response trained either concurrently or in separate components from the alternative response across conditions. Target response rates trained alone in baseline were higher but resistance to extinction and relapse via reinstatement tests were greater after training concurrently with the alternative response. In another assessment, training target and alternative responding together, but separating them during extinction and reinstatement tests, produced equal resistance to extinction and relapse. Together, these findings are consistent with behavioral momentum theory-operant response-reinforcer relations determined baseline response rates but Pavlovian stimulus-reinforcer relations established during training determined resistance to extinction and relapse. These findings imply that reinforcing alternative behavior to treat problem behavior could initially reduce rates but increase persistence.

Supplementation of complex milk lipid concentrate (CMLc) improved the memory of aged rats

Abstract Objectives The socio-economic impact from age-related mental decline is escalating. Supplementation of functional foods for sustaining mental health is desirable. We examined the effect of long-term supplementation of complex milk lipid concentrate (CMLc), mixed dairy phospholipids, on memory and associated vascular and neuronal changes in aged rats. Methods Fisher/Norway Brown rats were used. Two groups of aged rats (24 months) were fed with either gelatin-formulated CMLc or blank gelatin as the control, for 4 months. To determine age-related changes, a young group (5 months) was also fed with blank gelatin. Morris water maze tests were carried out after the supplementation and brain tissues were collected for biological analysis. Results The aged control rats learnt to locate the platform slower than the young control rats during acquisition trials (*P < 0.05), and made fewer entries to and more initial heading errors from the platform zone during testing trials (*P < 0.05). The CMLc supplementation improved memory by showing the reduced initial heading errors in a delayed probe trial (#P < 0.05). We also found that the aged rats with CMLc supplementation improved vascular density, dopamine output, and neuroplasticity (#P < 0.05) in the brain regions involved in memory compared with that of the aged control rats. Discussion The data suggested that the supplementation of CMLc during the early stage of brain aging may prevent memory decline possibly through improving vascular and neuronal function.

Choice, time and food: continuous cyclical changes in food probability between reinforcers.

The current experiment examined the degree to which locally varying food probabilities on two keys across time since food presentations can continue to control choice until the next food delivery. In two sets of conditions, the probability of food delivery being made available on one key relative to the other key varied sinusoidally across a 1-min period following each food delivery. In Set 1, food-probability changes were unsignaled and the number of cycles per min was varied across conditions. In Set 2, there were always two complete cycles of the sinusoid in the 1-min period, and brief key-color changes were arranged at a selection of fixed times since food delivery to signal portions of the sinusoid. In Set 1, control of choice by local probability of food on each key decreased over time since food delivery. Control by local food probabilities was greater in conditions that arranged fewer cycles per min. The onset of stimulus changes in Set 2 led to a transient reinstatement of local control by food probabilities regardless of the portion of the sinusoidal variation in food probabilities signaled by the stimuli. However, in conditions where the same colored stimuli signaled different portions of the sinusoidal variation in food-delivery probabilities, stimulus changes attenuated joint control by elapsed time and food-probability values. These results suggest that, changing relative food probabilities and stimuli can direct preference toward the likely location of the next food delivery across time since a food presentation, although the degree to which control over choice will be maintained across elapsed time depends on how experimenter-arranged contingencies are mapped onto elapsed time.