Wikipedia

Spontaneous alternation

Spontaneous Alternation Behavior (SAB) describes the tendency to alternate in the pursuit of different stimuli in consecutive trials, despite a lack of training or reinforcement.[1][2] The behavior emerged from experiments using animals, mainly rodents, who naturally demonstrated the behavioral pattern when placed in previously unexplored maze shapes (e.g. using a T/Y-maze).[2][3]

Spontaneous alternation testing is a behavioral assessment method derived from SAB. It is used to investigate exploratory behavior[4] and cognitive function (related to spatial learning and memory).[4][5] These assessments are most often done with non-human animals. The test serves great purpose in comparative psychology,[6] wherein non-human animals are studied to investigate differences within and between species with the aims of applying their findings to a greater understanding of human behavior.[7] It is particularly useful in studying the potential neuroanatomical and neurobiological mediators of cognitive function,[8] seeing that there are ethical limitations posed in the physiological study of humans, there is greater opportunity for more invasive procedures to be ethically conducted on non-human animals.

Explanations for spontaneous alternation

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Earlier explanations for SAB thought the behavior to be a result of reactive inhibition. According to Hull's concept, the animal turns right in a T-maze after it has already turned left, as turning left the first time diminished the value of the tendency to do so again.[2][8][9] The inhibitive response is cumulative and therefore repeats, resulting in SAB.[10] Several experiments conducted in the 1950s found reason to refute this theory, demonstrating SAB to be a stimulus-based behavior, as opposed to one which is response-based.[8][11] These experiments point towards perception, attention, memory and motivation as psychological processes which produce SAB.

Perception: The pursuit of directional movement relies on the subject's manner of gathering information from its surroundings.[8]

Attention: The subject selectively abstracts cues from the greater informational environment, informing their concentrate focus and influencing behavior.[8]

Memory: In order to demonstrate a consistent alternating pattern in behavior, the cues which are attended to and perceived from the first arm of the maze must be stored and retrieved for the pursuit of the alternate arm to be initiated.[8]

Motivation: As movement into either arm of the maze isn't traced, nor reinforced, the animal must have a purpose which is fulfilled by SAB that is not exclusively cognitive. Experimental research suggests that the behavior is underpinned by an innate drive towards exploring that which is unfamiliar, or less familiar than the other alternative/s.[8] Dember and Earl (1957) describe this to be 'a manifestation of exploratory motivation' within their general theory of exploration and curiosity.[2][8][12] The implied hardwiring of the behavior in the brain draws on the concept of evolutionary psychology. Estes and Schoeffler (1955) proposed that SAB originated as an adaptive behavior which enabled animals to gather information related to the distribution of resources needed for survival (e.g. food and shelter). With this logic, an animal is likely motivated to enter environments in an alternating pattern, because they will have already exhausted the resources from that which they most recently visited.[8]

Spontaneous alternation and spatial working memory

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There is a comprehensive amount of research which suggests that efficient spatial alternation behaviour requires optimal spatial working memory.[13]

Spatial working memory refers to the ability to retain spatial information, in the form of environmental cues, in working memory, where it is stored temporarily and elicited actively during the completion of a task.[14] This allows one to build, and continuously update, cognitive spatial maps of novel environments as they are being explored, so that they are able to return to areas which they distinguish as less familiar – thereby optimizing information gathering strategies.[13]

Therefore, in tests where SAB is reduced, there is an indication of impairment in spatial working memory, seeing as one's ability to distinguish where they have or have not been is diminished.[15] Hence, spontaneous alternation tests are useful in investigating spatial working memory and the factors which may influence it.

Aging was found to be a factor which influences SAB. Spontaneous alternation testing with a Y-maze found that the frequency of the behavior reduces with age in mice at 9- and 12- months of age.[16] Variation in physiological ability (e.g. motor function), did not account for this, instead, the observations were attributed to gradual degradation in spatial learning.[13][16] Anatomical changes to limbic and non-limbic pathways associated with normal and pathological aging in rodents correlate with the decrease in SAB. These include the neurochemical pathways in the hippocampus and basal forebrain– both of which are associated with spatial working memory and learning in humans, as well as in rodents. Legions in the brain (particularly in the hippocampus), and various drugs can significantly impair spontaneous alteration.[17] In another study done with mice, increasing the hormone estrogen showed improved spontaneous alteration performance.[18] These findings may suggest a decrease in exploratory behavior as a result of associated neurobiological changes, which implicate the capacity and function of spatial working memory.[19]

Apparatus design and test calculations

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The apparatus used for spontaneous alternation testing takes multiple forms – the T-maze and the Y-maze being those which are most commonly used in experimental psychology. Both apparatus are named to mimic the maze shapes which they portray.[2][4][8][20] The rat is placed in the middle of the maze, and is allowed to move freely through each arm. Variations in apparatus (e.g. shape, material, incline etc.), can result in subtle changes to the rate of SAB, though the general behavioral pattern persists.[2]

SAB can be evaluated as a percentage indicating how frequently spontaneous alternation is observed across several trials. There are three outputs of the spontaneous alternation test.[21] One output is the number of entries one makes into each arm of the maze – one entry being defined by when the hind paws of the subject species crosses the boundary distinguishing one arm completely. Another output is the number of alternations the animal makes – one alternation being the number of consecutive entries into each arm of the maze without any repeats in any order. These are then put into the following equation to yield percentage alternation:

 

The cognition of the animal can be assessed based on the score, with a lower score is considered cognitively impaired. Chance is determined by the number of arms the maze has, in a three-arm maze it is 22%, in a four-arm or greater maze, it is around 9%.

Human infants

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Spontaneous alteration was tested in human infants at the ages of 6 months and 18 months.[22] The young children were presented two identical toys placed in different spots.[22] The six-month-old infants had no significant interchange with which toy they chose in subsequent trials, so the pattern of spontaneous alternation was not present.[22] However, the 18-month-old infants did alternate between the toys, curiosity motivating them to try the toy they had not chosen (the novel toy), displaying spontaneous alternation as well as spatial memory.[22] Although both ages showed inhibition of return, which is described as reacting to an object that has not been seen before, only the 18-month-old children actually chose the novel toy.[22][23] This suggests that inhibitory control, or voluntary control over actions, is needed to, at least physically, perform spontaneous alteration.[22]

Stress

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When testing the effects of stress on spontaneous alteration in mice, two different methods of stressors have been involved in research studies.[24] The first is called the inescapable stressor, where the mouse could not escape the bright light being shined into the maze (open field test), and escapable, where the light was shining into the maze, but there were areas that were covered, and therefore were dark for the mice to find shelter.[24] These methods caused stress, because mice prefer dim areas to bright areas.[24] The inescapable stressor (the bright light that lit up the entire maze), caused the mice's spontaneous alteration to decrease, but the escapable stressor (where only some parts of the maze were lit up), did not have an effect on their levels of spontaneous alteration.[24]

Criticisms

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Animal Models: As a method predominantly used in studies involving non-human animals, there are inherent criticisms which arise from the extent to which findings can extend towards humans in comparative psychology.[25] While the spontaneous alternation test was designed to gain insight into human cognition and behavior, its direct replication using human subjects is rarely conducted. Life-sized mazes are impractical to create, and lack naturalism in their implementation, eliciting a number of confounding variables related to social desirability and demand characteristics. Proposals have been made for the use of virtual reality mazes to test SAB, with one study reporting that SAB was observed in a VR representation of a T-maze.[26]

The Mere-Exposure Hypothesis: Zajonc's (1968) hypothesis reflects the tendency for animals to be drawn towards stimuli which is familiar.[27] Addressing this contradiction to SAB, some experiments suggest that SAB is more likely to be demonstrated after the animal has been subjected to a period of long-term confinement to one environment, following which a preference for novel environments is built.[27]

References

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  1. ^ d'Isa, R.; Comi, G.; Leocani, L. (2021). "Apparatus design and behavioural testing protocol for the evaluation of spatial working memory in mice through the spontaneous alternation T-maze". Scientific Reports. 11 (1): 21177. Bibcode:2021NatSR..1121177D. doi:10.1038/s41598-021-00402-7. PMC 8551159. PMID 34707108.
  2. ^ a b c d e f Dember, W. N., & Richman, C. L. (2012). Spontaneous Atlernation Behaviour. Springer.
  3. ^ Dennis, W. (1935). A Comparison of the Rat's First and Second Explorations of a Maze Unit. The American Journal of Psychology, 47(3), 488. doi:10.2307/1416343
  4. ^ a b c T Maze Spontaneous Alternation. (n.d.). [TLD]. Stanford Medicine. Retrieved 22 March 2020
  5. ^ Wolf, A., Bauer, B., Abner, E. L., Ashkenazy-Frolinger, T., & Hartz, A. M. S. (2016). A Comprehensive Behavioral Test Battery to Assess Learning and Memory in 129S6/Tg2576 Mice. PLoS One, 11(1). doi:10.1371/journal.pone.0147733
  6. ^ The Editors of Encyclopaedia Britannica. (2020). Comparative psychology. In Encyclopædia Britannica. Encyclopædia Britannica, inc.
  7. ^ Domjan, M. (1987). Comparative Psychology and the Study of Animal Learning. Journal of Comparative Psychology, 101(3), 237–241. doi:10.1037/0735-7036.101.3.237
  8. ^ a b c d e f g h i j Richman, C. L., Dember, W. N., & Kim, P. (1986). Spontaneous Alternation Behavior in Animals: A Review. Current Psychological Research & Reviews, 5, 358–391. doi:10.1007/BF02686603
  9. ^ Ryan, R. M. (2012). The Oxford Handbook of Human Motivation. Oxford University Press.
  10. ^ Glanzer, M. (1953). Stimulus satiation: An explanation of spontaneous alternation and related phenomena. Psychological Review, 60(4), 257–268. doi:10.1037/h0062718
  11. ^ Wayne, D. (1939). Spontaneous alternation in rats as an indicator of the persistence of stimulus effects. Journal of Comparative Psychology, 28(2), 305–312. doi:10.1037/h0056494
  12. ^ Dember, W. N., & Earl, R. W. (1957). Analysis of exploratory, manipulatory, and curiosity behaviors. Psychological Review, 64(2), 91–96. doi:10.1037/h0046861
  13. ^ a b c Lewis, S. A., Negelspach, D. C., Kaladchibachi, S., Cowen, S. L., & Fernandez, F. (2017). Spontaneous alternation: A potential gateway to spatial working memory in Drosophila. Neurobiology of Learning and Memory, 142, 230–235. doi:10.1016/j.nlm.2017.05.013
  14. ^ van Asselen, M., Kessels, R. P., Neggers, S. F., Kappelle, L. J., Frijns, C. J., & Postma, A. (2005). Brain areas involved in spatial working memory. Neuropsychologia, 44(7), 1185–1194. doi:10.1016/j.neuropsychologia.2005.10.005
  15. ^ Li, B., Arime, Y., Hall, F. S., & Uhl, G. R. (2009). Impaired spatial working memory and decreased frontal cortex BDNF protein level in dopamine transporter knockout mice. European Journal of Pharmacology, 628(1–3), 104–107. doi:10.1016/j.ejphar.2009.11.036
  16. ^ a b Lamberty, Y., & Gower, A. J. (1990). Age-related changes in spontaneous behavior and learning in NMRI mice from maturity to middle age. Physiology & Behavior, 47(6), 1137–1144. doi:10.1016/0031-9384(90)90364-A
  17. ^ Douglas, Robert J. (1989), Dember, William N.; Richman, Charles L. (eds.), "Spontaneous Alternation Behavior and the Brain", Spontaneous Alternation Behavior, Springer, pp. 73–108, doi:10.1007/978-1-4613-8879-1_5, ISBN 978-1-4613-8879-1, retrieved 2020-03-17
  18. ^ Miller, M. M.; Hyder, S. M.; Assayag, R.; Panarella, S. R.; Tousignant, P.; Franklin, K. B. J. (1999-07-01). "Estrogen modulates spontaneous alternation and the cholinergic phenotype in the basal forebrain". Neuroscience. 91 (3): 1143–1153. doi:10.1016/S0306-4522(98)00690-3. ISSN 0306-4522.
  19. ^ Adelöf, J., Ross, M., Lazic, S., Zetterberg, M., Wiseman, J., & Hernebring, M. (2019). Conclusions from a behavioral aging study on male and female F2 hybrid mice on age-related behavior, buoyancy in water-based tests, and an ethical method to assess lifespan.Aging, 11(17), 7150–7168. doi:10.18632/aging.102242
  20. ^ Y Maze Spontaneous Alternation Test. (n.d.). [TLD]. Stanford Medicine. Retrieved 22 March 2020
  21. ^ Ohno, M., Sametsky, E. A., Younkin, L. H., Oakley, H., Younkin, S. G., Citron, M., Vassar, R., & Disterhoft, J. F. (2004). BACE1 deficiency rescues memory deficits and cholinergic dysfunction in a mouse model of Alzheimer's disease. Neuron, 41(1), 27–33. doi:10.1016/S0896-6273(03)00810-9
  22. ^ a b c d e f Vecera, Sham P.; Rothbart, Mary K.; Posner, Michael I. (1991-10-01). "Development of Spontaneous Alternation in Infancy". Journal of Cognitive Neuroscience. 3 (4): 351–354. doi:10.1162/jocn.1991.3.4.351. ISSN 0898-929X.
  23. ^ Dukewich, Kristie R.; Klein, Raymond M. (2015-07). "Inhibition of return: A phenomenon in search of a definition and a theoretical framework". Attention, Perception, & Psychophysics. 77 (5): 1647–1658. doi:10.3758/s13414-015-0835-3. ISSN 1943-3921.
  24. ^ a b c d Bats, S; Thoumas, J. L; Lordi, B; Tonon, M. C; Lalonde, R; Caston, J (2001-01-08). "The effects of a mild stressor on spontaneous alternation in mice". Behavioural Brain Research. 118 (1): 11–15. doi:10.1016/S0166-4328(00)00285-0. ISSN 0166-4328.
  25. ^ Bracken, M. B. (2008). Why animal studies are often poor predictors of human reactions to exposure. Journal of the Royal Society of Medicine, 102(3), 120–122. doi:10.1258/jrsm.2008.08k033
  26. ^ Rothacher, Y., Nguyen, A., Lenggenhager, B., Kunz, A., & Brugger, P. (2020). Walking through virtual mazes: Spontaneous alternation behaviour in human adults. Cortex, 127, 1–16. doi:10.1016/j.cortex.2020.01.018
  27. ^ a b Syme, G. J., & Syme, L. A. (1977). Spontaneous Alternation in Mice: A Test of the Mere-Exposure Hypothesis. The American Journal of Psychology, 90(4), 621–633. doi:10.2307/1421736
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