蜜蜂对复合气味中特征因素的选择性记忆巩固的研究

目的 蜜蜂天生具有丰富的嗅觉辨识能力,觅食、交配、导航以及社交活动均依赖其嗅觉系统,是研究嗅觉感知和学习记忆的行为及神经机制的理想模型。蜜蜂既能够将某个复合气味作为一个整体也可以将复合气味的各组成成分进行辨别和区分,但是在特征依赖的联合记忆中依据何种原则进行加工并存储到长期记忆还不清楚。方法 本文利用特征阳性（feature positive：AB+,B-）和特征阴性（feature negative：AB-,B+）的奖赏性嗅觉条件化,训练蜜蜂对复合气味和成分气味的辨别,并检测蜜蜂对复合气味（AB）、成分气味（B）以及特征气味（A）的中长时记忆（3 h）和长时记忆（24 h）。结果 在特征阳性的奖赏性嗅觉条件化中,蜜蜂对训练过的气味可以形成稳定的中长时和长时记忆,并且对复合气味中的特征气味的记忆与复合气味的记忆呈现高度相似。但在特征阴性的奖赏性嗅觉条件化中,蜜蜂虽能够在3 h和24 h对训练过的两种气味具有显著的伸喙反应差异,且对特征阴性的气味无显著反应,但对复合气味的反应随时间的推移而增加。结论 实验结果表明,蜜蜂选择性地将与奖赏信息联合出现的气味巩固到长时记忆中,但并未依据特征成分加工储存到长时记忆中。奖赏信息预示着食物源,与生存息息相关,表明对环境信息进行选择性的记忆巩固加工并储存可能是低等动物高效地编码生存相关信息的重要策略。     

Input coding in animal and human associative learning

Two appetitive conditioning experiments with rats investigated whether the degree of generalization between a compound and its component parts is fixed or variable. Both experiments used a two-stage transfer design. In Stage 1, the elemental groups learned that a compound and its component parts signaled the same outcome (i.e. C+, D+, CD+), whereas the configural groups learned that a compound and its component parts signaled different outcomes (i.e. C+, D+, CD-, where '+' is pellets and '-' is no pellets). In Stage 2, the rats were tested for reductions in generalization. Experiment 1 found no evidence that past configural learning reduced generalization when a new set of alike-treated A and B elements were presented in compound for the first time. Experiment 2 found no evidence that past configural learning reduced generalization when the stimuli of Stage 1 were presented in a new C-, D-, CD+ relation. In contrast to findings with humans, these results suggest that past experience plays a minor role in how stimuli are encoded in animal conditioning.     

Indicators of Early and Late Processing Reveal the Importance of Within-Trial-Time for Theories of Associative Learning

In four human learning experiments (Pavlovian skin conductance, causal learning, speeded classification task), we evaluated several associative learning theories that assume either an elemental (modified unique cue model and Harris’ model) or a configural (Pearce’s configural theory and an extension of it) form of stimulus processing. The experiments used two modified patterning problems (A/B/C+, AB/BC/AC+ vs. ABC-; A+, BC+ vs. ABC-). Pearce’s configural theory successfully predicted all of our data reflecting early stimulus processing, while the predictions of the elemental theories were in accord with all of our data reflecting later stages of stimulus processing. Our results suggest that the form of stimulus representation depends on the amount of time available for stimulus processing. Our findings highlight the necessity to investigate stimulus processing during conditioning on a finer time scale than usually done in contemporary research.     

SSCC TD: A Serial and Simultaneous Configural-Cue Compound Stimuli Representation for Temporal Difference Learning

This paper presents a novel representational framework for the Temporal Difference (TD) model of learning, which allows the computation of configural stimuli – cumulative compounds of stimuli that generate perceptual emergents known as configural cues. This Simultaneous and Serial Configural-cue Compound Stimuli Temporal Difference model (SSCC TD) can model both simultaneous and serial stimulus compounds, as well as compounds including the experimental context. This modification significantly broadens the range of phenomena which the TD paradigm can explain, and allows it to predict phenomena which traditional TD solutions cannot, particularly effects that depend on compound stimuli functioning as a whole, such as pattern learning and serial structural discriminations, and context-related effects.     

Nonelemental visual learning in honeybees

Free-flying honeybees, Apis mellifera, learn visual stimuli in the appetitive context of food search. Visual compound stimuli are relevant in nautre as bees learn flower images that consist of many visual elements. We studied whether elemental associations between each visual element and the reinforcement (elemental approach) are enough to explain the solving of visual discrimination problems that raise ambiguity at the elemental level. We asked whether bees could solve three different visual discriminations: (1) positive patterning (A−, B−, AB+); (2) negative patterning (A+, B+, AB−); and (3) biconditional discrimination (AB+, CD+, AC−, BD−). In experiments 1 and 2 bees had to discriminate a yellow-violet chequerboard from the yellow or the violet squares alone. In experiment 3, four different gratings combining one colour (yellow or violet) with one orientation (vertical or horizontal) had to be discriminated. In all three problems binary compounds were trained in such a way that each element appeared equally often as rewarded and nonrewarded. Bees could solve the three discrimination problems. They always chose the reinforced stimulus despite ambiguity at the level of the elements. For solving positive patterning, elemental processing could be used. For negative patterning and biconditional discrimination, nonelemental processing strategies (unique-cue or configural approach) are necessary to account for these results. Although we cannot decide between a configural and a unique-cue interpretation, we can clearly reject purely elemental processing in these cases. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Effects of atropine on operant test battery performance in rhesus monkeys.

The acute behavioral effects of atropine sulfate were assessed using a battery of complex food-reinforced operant tasks that included: temporal response differentiation (TRD, n = 7); delayed matching-to-sample (DMTS, n = 6), progressive ratio (PR, n = 8), incremental repeated acquisition (IRA, n = 8), and conditioned position responding (CPR, n = 8). Performance in these tasks is thought to depend primarily upon specific brain functions such as time perception, short-term memory and attention, motivation, learning, and color and position discrimination, respectively. Atropine sulfate (0.01-0.56 mg/kg iv), given 15-min pretesting, produced significant dose-dependent decreases in the number of reinforcers obtained in all tasks. Response rates decreased significantly at greater than or equal to 0.03 mg/kg for the learning and discrimination tasks, at greater than or equal to 0.10 mg/kg for the motivation and short-term memory and attention tasks, and at greater than or equal to 0.30 mg/kg for the time perception task. Response accuracies were significantly decreased at doses greater than or equal to 0.10 mg/kg for the learning, discrimination, and short-term memory and attention tasks, and at greater than or equal to 0.30 mg/kg for the time perception task. Thus, the order of task sensitivity to any disruption by atropine is learning = color and position discrimination greater than time perception = short-term memory and attention = motivation (IRA = CPR greater than TRD = DMTS = PR). Thus in monkeys, the rates of responding in operant tasks designed to model learning and color and position discrimination were the most sensitive measures to atropine's behavioral effects. Accuracy in these same task was also disrupted but at higher doses. These data support the hypothesis that cholinergic systems play a greater role in the speed (but not accuracy) of performance of our learning and discrimination tasks compared to all other tasks. Accuracy of responding in these and the short-term memory task, all of which involve the use of lights as visual stimuli, was more sensitive to disruption by atropine than those tasks which did not utilize such strong visual stimuli.     

Occasion setting in Pavlovian ambiguous target discriminations

Rats were trained in a Pavlovian serial ambiguous target discrimination, in which a target cue was reinforced if it was preceded by one stimulus (P-->T+) but was not reinforced if it was preceded by another stimulus (N-->T-). Test performance indicated that stimulus control by these features was weaker than that acquired by features trained within separate serial feature positive (P-->T+, T-) and serial feature negative (N-->W-, W+) discriminations. The form of conditioned responding and the patterns of transfer observed suggested that the serial ambiguous target discrimination was solved by occasion setting. The data are discussed in terms of the use of retrospective coding strategies when solving Pavlovian serial conditional discriminations, and the acquisition of special properties by both feature and target stimuli.     

An examination of the effectiveness of inflation and deflation treatments in detecting within-compound learning of a taste aversion

Two conditioned taste aversion experiments with rats assessed the relative effectiveness in providing evidence of within-compound learning of different procedures that involve the initial compound presentation of two stimuli, A and X, with the unconditioned stimulus (i.e., AX+). In Experiment 1, following a single AX+ trial, groups A+ and B+ received an additional conditioning trial (i.e., inflation treatment) with A and B, respectively, whereas group A- received an extinction trial (i.e., deflation treatment) with A. The results showed a reduction in the aversion elicited by the target stimulus, X, in group A- relative to both groups A+ and B+, which did not differ. Experiment 2 further investigated the failure of group A+ to increase the aversion to X relative to control group B+ by pairing A or B with either the same unconditioned stimulus that was previously paired with AX (groups A+ and B+) or with a stronger unconditioned stimulus (groups A* and B*). The results showed increased aversion to X in group A* relative to group B*, but not in group A+ relative to group B+. These results are interpreted as indicative of extinction of the within-compound association during the treatment with A, which could likely impair the detection of within-compound learning following an inflation, but not a deflation treatment.     

Are There Age-Related Differences in the Ability to Learn Configural Responses?

Age is often associated with a decline in cognitive abilities that are important for maintaining functional independence, such as learning new skills. Many forms of motor learning appear to be relatively well preserved with age, while learning tasks that involve associative binding tend to be negatively affected. The current study aimed to determine whether age differences exist on a configural response learning task, which includes aspects of motor learning and associative binding. Young (M = 24 years) and older adults (M = 66.5 years) completed a modified version of a configural learning task. Given the requirement of associative binding in the configural relationships between responses, we predicted older adults would show significantly less learning than young adults. Older adults demonstrated lower performance (slower reaction time and lower accuracy). However, contrary to our prediction, older adults showed similar rates of learning as indexed by a configural learning score compared to young adults. These results suggest that the ability to acquire knowledge incidentally about configural response relationships is largely unaffected by cognitive aging. The configural response learning task provides insight into the task demands that constrain learning abilities in older adults.     

Baboons (Papio papio) spontaneously process the first-order but not second-order configural properties of faces

A two-alternative forced-choice discrimination task was used to assess whether baboons (N=7) spontaneously process qualitative (i.e., first-order) or quantitative (i.e., second-order) variations in the configural arrangement of facial features. Experiment 1 used as test stimuli second-order pictorial faces of humans or baboons in which the mouth and the eyes were rotated upside down relative to the normal face. Baboons readily discriminated two different normal faces but did not discriminate a normal face from its second-order modified version. Experiment 2 used human or baboon faces for which the first-order configural properties had been distorted by reversing the location of the eyes and mouth within the face. Discrimination was prompt with these stimuli. Experiment 3 replicated some of the conditions and the results of experiment 1, thus ruling out possible effects of learning. It is concluded that baboons are more adept at spontaneously processing first- than second-order configural facial properties, similar to what is known in the human developmental literature.     

US specificity of occasion setting: Hierarchical or configural learning?

Four experiments in rats examined whether occasion setters and target CSs play qualitatively different roles in occasion-setting discriminations. Two visual occasion setters, A and B, signalled reinforcement of two auditory target CSs, x and y, with sucrose and oil (A…x→suc, B…y→oil, A-, B-, x-, y-); in addition two transfer CSs w and z were paired with sucrose and oil (w→suc, z→oil). When w and z were substituted for x and y (A…w, B…w, A…z, B…z) more responding was observed when both stimuli had been paired with the same outcome (Experiments 1 and 3a). No effect was observed when two visual "pseudo-occasion setters", C and D (paired with sucrose and oil in a trace relation to the US:C…→suc, D…→oil), were substituted for the occasion setters A and B (C…x, D…x, C…y, D…y; Experiments 2, 3b and 4). These results could not be explained in terms of Pavlovian summation: responding to combinations of Pavlovian CSs paired with same or different outcomes was either the same, or lower when both stimuli had been paired with the same outcome (Experiment 4). Implications of these results for theories of occasion setting and configural learning are discussed.     

Transient Decline in Hippocampal Theta Activity during the Acquisition Process of the Negative Patterning Task

Hippocampal function is important in the acquisition of negative patterning but not of simple discrimination. This study examined rat hippocampal theta activity during the acquisition stages (early, middle, and late) of the negative patterning task (A+, B+, AB-). The results showed that hippocampal theta activity began to decline transiently (for 500 ms after non-reinforced stimulus presentation) during the late stage of learning in the negative patterning task. In addition, this transient decline in hippocampal theta activity in the late stage was lower in the negative patterning task than in the simple discrimination task. This transient decline during the late stage of task acquisition may be related to a learning process distinctive of the negative patterning task but not the simple discrimination task. We propose that the transient decline of hippocampal theta activity reflects inhibitory learning and/or response inhibition after the presentation of a compound stimulus specific to the negative patterning task.     

与作业无关的新异刺激对猕猴额叶神经元延缓期反应的作用

在猕猴执行延缓辨别作业和单纯辨别作业时,观察了与作业无关的新异刺激对额叶神经元延缓期放电的影响。在这两种作业中,延缓期在1—4s之间随机变化。此时,动物必须高度注意信号的变化,稍不注意即导致操作错误。此外,在延缓辨别作业中,动物在延缓期还要暂时记住暗示期的信号,单纯辨别作业则无此要求。在203个与作业相关的神经元中,有70个神经元在延缓期出现放电频率变化,其中见于延缓辨别作业者41个,见于单纯辨别作业者29个。实验结果表明,在这两种作业的延缓期所出现的神经元放电增多的反应,有着许多相同的特点。与课题无关的声、光、触、痛等刺激引起分心时,神经元的延缓期反应出现明显的变化,随之出现操作错误。多数神经元的反应受到抑制,但也有出现反应增强者,而且同一神经元对不同感觉模式的无关刺激可出现不同的效应,表现出不同程度的感觉模式特异性。此外,无关刺激在延缓期和在测试间歇期可产生不同甚至相反的效应。上述在延缓期出现反应的神经元主要位于额叶弓状沟上支内侧部的一定范围内。本文对实验结果进行了讨论,认为额叶神经元的延缓期反应,可能在很大程度上与注意有关。额叶神经元感觉模式各种程度的特异性可能是注意的通道选择性的神经基础。额叶的背内侧部,包括前额叶后部和运动前区前部     

The Categorisation of Non-Categorical Colours: A Novel Paradigm in Colour Perception

In this paper, we investigate a new paradigm for studying the development of the colour ‘signal’ by having observers discriminate and categorize the same set of controlled and calibrated cardinal coloured stimuli. Notably, in both tasks, each observer was free to decide whether two pairs of colors were the same or belonged to the same category. The use of the same stimulus set for both tasks provides, we argue, an incremental behavioural measure of colour processing from detection through discrimination to categorisation. The measured data spaces are different for the two tasks, and furthermore the categorisation data is unique to each observer. In addition, we develop a model which assumes that the principal difference between the tasks is the degree of similarity between the stimuli which has different constraints for the categorisation task compared to the discrimination task. This approach not only makes sense of the current (and associated) data but links the processes of discrimination and categorisation in a novel way and, by implication, expands upon the previous research linking categorisation to other tasks not limited to colour perception.     

Different neuroplasticity for task targets and distractors

Adult learning-induced sensory cortex plasticity results in enhanced action potential rates in neurons that have the most relevant information for the task, or those that respond strongly to one sensory stimulus but weakly to its comparison stimulus. Current theories suggest this plasticity is caused when target stimulus evoked activity is enhanced by reward signals from neuromodulatory nuclei. Prior work has found evidence suggestive of nonselective enhancement of neural responses, and suppression of responses to task distractors, but the differences in these effects between detection and discrimination have not been directly tested. Using cortical implants, we defined physiological responses in macaque somatosensory cortex during serial, matched, detection and discrimination tasks. Nonselective increases in neural responsiveness were observed during detection learning. Suppression of responses to task distractors was observed during discrimination learning, and this suppression was specific to cortical locations that sampled responses to the task distractor before learning. Changes in receptive field size were measured as the area of skin that had a significant response to a constant magnitude stimulus, and these areal changes paralleled changes in responsiveness. From before detection learning until after discrimination learning, the enduring changes were selective suppression of cortical locations responsive to task distractors, and nonselective enhancement of responsiveness at cortical locations selective for target and control skin sites. A comparison of observations in prior studies with the observed plasticity effects suggests that the non-selective response enhancement and selective suppression suffice to explain known plasticity phenomena in simple spatial tasks. This work suggests that differential responsiveness to task targets and distractors in primary sensory cortex for a simple spatial detection and discrimination task arise from nonselective increases in response over a broad cortical locus that includes the representation of the task target, and selective suppression of responses to the task distractor within this locus.     

Auditory Discrimination Learning: Role of Working Memory

Perceptual training is generally assumed to improve perception by modifying the encoding or decoding of sensory information. However, this assumption is incompatible with recent demonstrations that transfer of learning can be enhanced by across-trial variation of training stimuli or task. Here we present three lines of evidence from healthy adults in support of the idea that the enhanced transfer of auditory discrimination learning is mediated by working memory (WM). First, the ability to discriminate small differences in tone frequency or duration was correlated with WM measured with a tone n-back task. Second, training frequency discrimination around a variable frequency transferred to and from WM learning, but training around a fixed frequency did not. The transfer of learning in both directions was correlated with a reduction of the influence of stimulus variation in the discrimination task, linking WM and its improvement to across-trial stimulus interaction in auditory discrimination. Third, while WM training transferred broadly to other WM and auditory discrimination tasks, variable-frequency training on duration discrimination did not improve WM, indicating that stimulus variation challenges and trains WM only if the task demands stimulus updating in the varied dimension. The results provide empirical evidence as well as a theoretic framework for interactions between cognitive and sensory plasticity during perceptual experience.     

Serial order of conditional stimuli as a discriminative cue for Pavlovian conditioning

The serial order in which events occur can be a signal for different outcomes and therefore might be a determinant of how an animal should respond. In this report, we propose a novel design for studying serial order learning in Pavlovian conditioning. In both Experiments 1a and 1b, hungry rats were trained with successively presented pairs of auditory and visual stimuli (e.g., A --> B) using four different stimuli (A-D). Four orders were paired with food (A --> B, B --> C, C --> D, D --> A) while the reversals were extinguished (B --> A, C --> B, D --> C, A --> D). An analysis of responding from the second element of each pair showed that the rats discriminated trial types that preceded food from those that did not. A replication of the effect using a completely counterbalanced design is described in Experiment 1b. These results suggest that rats can use the serial or temporal order of two sequentially presented non-overlapping elements as the basis for discrimination. Two associative accounts are suggested as possible mechanisms for solving the discrimination.     

Does Presentation Format Influence Visual Size Discrimination in Tufted Capuchin Monkeys (Sapajus spp.)?

Most experimental paradigms to study visual cognition in humans and non-human species are based on discrimination tasks involving the choice between two or more visual stimuli. To this end, different types of stimuli and procedures for stimuli presentation are used, which highlights the necessity to compare data obtained with different methods. The present study assessed whether, and to what extent, capuchin monkeys’ ability to solve a size discrimination problem is influenced by the type of procedure used to present the problem. Capuchins’ ability to generalise knowledge across different tasks was also evaluated. We trained eight adult tufted capuchin monkeys to select the larger of two stimuli of the same shape and different sizes by using pairs of food items (Experiment 1), computer images (Experiment 1) and objects (Experiment 2). Our results indicated that monkeys achieved the learning criterion faster with food stimuli compared to both images and objects. They also required consistently fewer trials with objects than with images. Moreover, female capuchins had higher levels of acquisition accuracy with food stimuli than with images. Finally, capuchins did not immediately transfer the solution of the problem acquired in one task condition to the other conditions. Overall, these findings suggest that – even in relatively simple visual discrimination problems where a single perceptual dimension (i.e., size) has to be judged – learning speed strongly depends on the mode of presentation.     

Does Perceptual Learning Suffer from Retrograde Interference?

In motor learning, training a task B can disrupt improvements of performance of a previously learned task A, indicating that learning needs consolidation. An influential study suggested that this is the case also for visual perceptual learning. Using the same paradigm, we failed to reproduce these results. Further experiments with bisection stimuli also showed no retrograde disruption from task B on task A. Hence, for the tasks tested here, perceptual learning does not suffer from retrograde interference.     

Towards an understanding of the mechanisms of weak central coherence effects: experiments in visual configural learning and auditory perception

The weak central coherence hypothesis of Frith is one of the most prominent theories concerning the abnormal performance of individuals with autism on tasks that involve local and global processing. Individuals with autism often outperform matched nonautistic individuals on tasks in which success depends upon processing of local features, and underperform on tasks that require global processing. We review those studies that have been unable to identify the locus of the mechanisms that may be responsible for weak central coherence effects and those that show that local processing is enhanced in autism but not at the expense of global processing. In the light of these studies, we propose that the mechanisms which can give rise to 'weak central coherence' effects may be perceptual. More specifically, we propose that perception operates to enhance the representation of individual perceptual features but that this does not impact adversely on representations that involve integration of features. This proposal was supported in the two experiments we report on configural and feature discrimination learning in high-functioning children with autism. We also examined processes of perception directly, in an auditory filtering task which measured the width of auditory filters in individuals with autism and found that the width of auditory filters in autism were abnormally broad. We consider the implications of these findings for perceptual theories of the mechanisms underpinning weak central coherence effects.