Richard Gerrit Coss is an American evolutionary psychologist and academic. He is a Professor Emeritus of Psychology at the University of California, Davis.[1]

Richard Gerrit Coss
NationalityAmerican
Occupation(s)Evolutionary psychologist and academic
Academic background
EducationB.S. in Architecture
M.A. in Design
PhD in Comparative psychology
Alma materUniversity of Southern California
University of California
University of Reading
Academic work
InstitutionsUniversity of California

Coss's research interest spans the field of behavioral evolution, with a particular focus on analyzing adaptive variation in antipredator behavior in different populations, and the impact of developmental, physiological, and neurobiological constraints on behavior. He is most known for his work on dendritic spine plasticity, and early contributions to the field of evolutionary aesthetic preferences.[2][3] He has also authored and co-authored more than 130 peer-reviewed journal articles[4] and is the editor of the book Environmental Awareness: Evolutionary, Aesthetic and Social Perspectives.[5]

Coss is a Fellow of the Association for Psychological Science[6] and has been a member of numerous professional societies, including the Animal Behavior Society and the International Society for the Arts, Sciences and Technology.[7]

Early life and education edit

Coss was born on January 3, 1940, in Sanger, California. He is the son of Dr. Joe Glenn and Cornelia Geraldine Coss. After completing his early education, he enrolled at the University of Southern California and graduated in 1962 with a major in Industrial Design in the School of Architecture. Later in 1966, he earned his master's degree in design from the University of California, Los Angeles. In 1973, he completed his Ph.D. in Comparative psychology at the University of Reading in the UK, where Corinne Hutt was his dissertation adviser. For his dissertation, Coss conducted comparative research on the perceptual aspects of eye-like schemata in African jewelfish, mouse lemurs,[8] and autistic and typically developing children.[9]

Career edit

Coss worked at Douglas Aircraft Company from 1962 to 1966. Afterwards, in 1966 he held an appointment as Research Director at the Compagnie de l'Esthetique Industrielle in France that developed the corporate identity for the Shell Oil Corporation.[10] Later, in 1986, he received a Fellowship by the NASA-Ames Research Center where he aided in evaluating ways to enhance space-station interiors for long-term habitation.[11]

Coss started his academic career in 1971, as a lecturer of design at the University of California, Los Angeles. After earning his Ph.D. in psychology, he became an assistant professor of psychology at the University of California, Davis. He was later promoted to associate professor in 1978 and became a professor of psychology in 1984. Since 2014, he has been serving as Professor Emeritus of Psychology at the University of California, Davis.[1]

Research edit

At the beginning of his career, Coss focused his research on human-factors engineering with aerospace applications at Douglas Aircraft Company. In this context, he was appointed Project Engineer for a lunar-base design proposal to NASA using, for habitation, the spherical tanks manufactured for the second stage of the Saturn 5 lunar rocket. He has studied environmental aesthetics with respect to art and design. Later, his work was aimed at analyzing the antipredator behavior of several species in both field and laboratory conditions as model systems for understanding the development of innate behavior and aesthetic preferences in the context of human evolutionary history. He has published numerous articles in scientific journals and was the recipient of a patent for creating an apparatus for measuring pupillary dilation.[12]

Behavioral evolution and environmental aesthetics edit

 
Painting by Richard Coss (1981) derived from video showing a female California ground squirrel jumping back after being struck in the face by a rattlesnake. The low facial swelling observed afterwards prompted research on adaptive variation in ground squirrel physiological resistance to rattlesnake venom.

Coss published a monograph in 1965 that described his visual perception research based on his theory that human ancestors were the prey of predators for a sufficient evolutionary time to engender innate recognition of predator features including two-facing eyes, sharp teeth, and claws.[13] Subsequently, he posited that recognizing such specific provocative shapes enhanced emotional arousal in a manner that have had an impact on works of art,[14] architecture, and product design.[15][16]

While working on behavioral evolution, Coss identified that, despite having relaxed natural selection from snake predation for more than 300,000 years,[17] California ground squirrels are still capable of distinguishing their rattlesnake and gopher snake predators[18] as compared with Arctic ground squirrels that have lost this ability over 3 million years.[19] However, both species under prolonged relaxed selection have also lost their physiological resistance to rattlesnake venom.[20][21] Following this research, he hypothesized that "evolved cognitive behavior in humans might persist for as long as 3 million years of relaxed selection" and to test this hypothesis he assessed the preschool children's selection of schematic trees as a form of refuge from predators and their overall refuge-seeking behavior on the playground,[22] particularly by investigating whether historical sexual-size dimorphism plays a role in influencing such behavior.[23] Subsequently, he examined the remembrances in pre-school children and adults of where "something scary" was located in their imagination relative to their beds at night[24][25] producing findings which reinforced his argument that ancestral sources of natural selection might continue to bias modern aesthetic expression.[15]

Innate pattern recognition edit

Coss has been acknowledged for his substantial work on the innate pattern-recognition ability of different species. He elucidated that the salient effects of glossy and sparkling surface finishes attract infants and toddlers,[26] increasing the possibilities of endangering their life by drowning or suffocation from plastic bags[27] and described that viewing water has calming effects on adults.[28] Furthermore, he discovered that wild California ground squirrels,[29] white-faced capuchin monkeys,[30] and bonnet macaques are capable of recognizing their snake predators[31] by their size and scale patterns and, for bonnet macaques, their leopard predators by the spots on their coat.[32][33] He later documented that young human infants are visually attracted reliably to snake-scale and leopard-spot patterns.[34]

Dendritic spine plasticity edit

Coss evaluated the provocative effects of two-facing eyes in humans and worked on the brain development[35] and behavior[36] of jewel fish which led him to a cover article in Science.[37] His joint study with A. Globus revealed that, due to social deprivation, the formation of dendritic branches in the optic tectum was arrested as was the experience-based shortening of dendritic spine stems.[38] Moreover, a single threatening experience shortened dendritic spine stems in the optic tectum,[39][40] and enhanced fish behavioral excitability in a manner analogous to PTSD in humans. In related research, he examined developing honeybees and observed the shortening of dendritic spine stems on calycal interneurons with progressive nursing and foraging experiences[41] and also during their first orientation flight.[42]

Evolutionary constraints edit

Coss also conducted a research series to explore the sources of natural selection mediating human brain evolution. This led to experiments measuring flight distances of wild horses in Arizona and African zebras to an approaching human.[43] The lower fear of wild horses compared with zebras led to his hypothesis that extensive human hunting in Africa might have led to an arm's race for more competent hunting by humans to counter the increasingly evasive ability of wary prey.[44] Such an arm's race for enhanced visual and spatial competence in hunting might explain the enlargement of the human parietal cortex for accurate spear throwing as well the ability of modern humans to translate mental images into figurative art with significant cultural impact. In contrast, the lack of figurative art among European Neanderthals, who likely hunted less wary game with thrusting spears during the Middle Paleolithic, might account for the smaller size of the Neanderthal parietal cortex that limited their artistic expression.[45]

Animal welfare edit

Apart from working on environmental aesthetics and behavioral evolution, Coss has contributed to animal welfare by examining the impacts of human intrusions in wildlife habitats. He advised graduate students studying African wild dogs and found that black-tailed deer in coastal California unaccustomed to humans were highly cautious when a human approached.[46][47] While working in India, he studied the sleeping-site selection of bonnet macaques living near human settlements,[48][49] and also examined the decline in tigers caused by the loss of palatable vegetation for large tiger prey.[50] Also in India, he studied crop-raiding by Asian Elephants,[51] and found that crop-raiding could be thwarted when elephant-activated tiger growls were played back to them.[52][53] Another facet of his research on animal welfare involved the effects of crowding on brain development. Jewel fish crowded during development continue to school like younger fish[54] and their tectal interneurons look like those of younger fish, an indication that crowding restricts behavioral flexibility similar to that of developmental deprivation.[55][56]

Bibliography edit

Books edit

  • Mood Provoking Visual Stimuli: Their Origins and Applications (1965)
  • Environmental Awareness: Evolutionary, Aesthetic & Social Perspectives (2005) ISBN 978-0757520112

Selected articles edit

  • Coss, R. G. (1968). The ethological command in art. Leonardo 1, 273–287. The Ethological Command in Art
  • Coss, R. G. & Perkel, D. H. (1985). The function of dendritic spines: A review of theoretical issues. Behavioral and Neural Biology, 44, 151–185. The function of dendritic spines: A review of theoretical issues
  • Coss, R. G. (1999). Effects of relaxed natural selection on the evolution of behavior. Geographic Variation in Behavior: Perspectives on Evolutionary Mechanisms (Foster, S. & Endler, J. A., eds). Oxford University Press, Oxford, 180–208.
  • Coss, R. G. (2003). The role of evolved perceptual biases in art and design. Evolutionary Aesthetics (Voland, E. & Grammer, K (eds). Springer-Verlag, Heidelberg, 69–130. Evolutionary Aesthetics
  • Coss, R. G. (2017). Drawings of representational images by Upper Paleolithic humans and their absence in Neanderthals reflect historical differences in hunting wary game. Evolutionary Studies in Imaginative Culture, 1(2), 15–38. [1]
  • Coss, R. G., and Keller, C. M. (2022). Transient decreases in blood pressure and heart rate with increased subjective level of relaxation while viewing water compared with adjacent ground. Journal of Environmental Psychology, 81, 101794. Transient decreases in blood pressure and heart rate with increased subjective level of relaxation while viewing water compared with adjacent ground

References edit

  1. ^ a b "Richard Coss — People in the Social Science Departments at UC Davis". psychology.ucdavis.edu.
  2. ^ Coss, Richard G.; Towers, Steven R. (January 1, 1990). "Provocative Aspects of Pictures of Animals in Confined Settings". Anthrozoös. 3 (3): 162–170. doi:10.2752/089279390787057586 – via Taylor and Francis+NEJM.
  3. ^ Clearwater, Yvonne A.; Coss, Richard G. (March 9, 1991). "Functional Esthetics to Enhance Weil-Being in Isolated and Confined Settings". In Harrison, Albert A.; Clearwater, Yvonne A.; McKay, Christopher P. (eds.). From Antarctica to Outer Space. Springer. pp. 331–348. doi:10.1007/978-1-4612-3012-0_31. ISBN 978-1-4612-7759-0 – via Springer Link.
  4. ^ "Richard Coss". scholar.google.com.
  5. ^ Coss, Richard G. (March 9, 2005). Environmental awareness: evolutionary, aesthetic, and social perspectives. Kendall/Hunt Publishing Company. ISBN 9780757520112 – via Google Books.
  6. ^ "APS Fellows". member.psychologicalscience.org.
  7. ^ "Richard Coss".
  8. ^ Coss, Richard G. (1978). "Perceptual Determinants of Gaze Aversion by the Lesser Mouse Lemur (Microcebus murinus), the Role of Two Facing Eyes". Behaviour. 64 (3/4): 248–270. doi:10.1163/156853978X00053. JSTOR 4533873 – via JSTOR.
  9. ^ Coss, Richard G. (1979). "Perceptual Determinants of Gaze Aversion by Normal and Psychotic Children: The Role of Two Facing Eyes". Behaviour. 69 (3/4): 228–254. doi:10.1163/156853979X00494. JSTOR 4533974. PMID 508240 – via JSTOR.
  10. ^ "The Shell project of the Compagnie de l'Esthétique Industrielle in 1968: review and lessons from the unsuccessful project Didier Pacoud".
  11. ^ Barbour, Christopher G.; Coss, Richard G. (December 9, 1988). "Differential Color Brightness as a Body Orientation Cue". Human Factors: The Journal of the Human Factors and Ergonomics Society. 30 (6): 713–717. doi:10.1177/001872088803000607. PMID 3235088. S2CID 219975551.
  12. ^ "Apparatus for measuring the dilation of the pupil".
  13. ^ "Mood Provoking Visual Stimuli: Their Origins and Applications".
  14. ^ Coss, Richard G. (1968). "The Ethological Command in Art". Leonardo. 1 (3): 273–287. doi:10.2307/1571871. JSTOR 1571871. S2CID 191393295 – via JSTOR.
  15. ^ a b Coss, Richard G. (March 9, 2003). Voland, Eckart; Grammer, Karl (eds.). Evolutionary Aesthetics. Springer. pp. 69–130. doi:10.1007/978-3-662-07142-7_4 – via Springer Link.
  16. ^ Coss, Richard G. (March 9, 2020). Carroll, Joseph; Clasen, Mathias; Jonsson, Emelie (eds.). Evolutionary Perspectives on Imaginative Culture. Springer International Publishing. pp. 185–212. doi:10.1007/978-3-030-46190-4_10. S2CID 226718107 – via Springer Link.
  17. ^ Thompson, N. S. (May 31, 1995). Perspectives in Ethology: Volume 11: Behavioral Design. Springer Science & Business Media. ISBN 9780306449062 – via Google Books.
  18. ^ "Individual Variation in the Antisnake Behavior of California Ground Squirrels (Spermophilus beecheyi)".
  19. ^ Ronald O. Goldthwaite; Coss, Richard G.; Owings, Donald H. (1990). "Evolutionary Dissipation of an Antisnake System: Differential Behavior by California and Arctic Ground Squirrels in Above- and Below-Ground Contexts". Behaviour. 112 (3/4): 246–269. doi:10.1163/156853990X00220. JSTOR 4534840 – via JSTOR.
  20. ^ Poran, Naomie S.; Coss, Richard G.; Benjamini, Eli (January 1, 1987). "Resistance of California ground squirrels (Spermophilus Beecheyi) to the venom of the northern Pacific rattlesnake (Crotalus Viridis Oreganus): A study of adaptive variation". Toxicon. 25 (7): 767–777. doi:10.1016/0041-0101(87)90127-9. PMID 3672545 – via ScienceDirect.
  21. ^ Coss, Richard G.; Kevin L. Gusé; Poran, Naomie S.; Smith, David G. (1993). "Development of Antisnake Defenses in California Ground Squirrels (Spermophilus beecheyi): II. Microevolutionary Effects of Relaxed Selection from Rattlesnakes". Behaviour. 124 (1/2): 137–164. doi:10.1163/156853993X00542. JSTOR 4535086 – via JSTOR.
  22. ^ Coss, Richard G.; Moore, Michael (October 1, 2002). "Precocious Knowledge of Trees as Antipredator Refuge in Preschool Children: An Examination of Aesthetics, Attributive Judgments, and Relic Sexual Dinichism". Ecological Psychology. 14 (4): 181–222. doi:10.1207/S15326969ECO1404_1. S2CID 145653776 – via Taylor and Francis+NEJM.
  23. ^ Coss, Richard G. (July 1, 2016). "Sex difference in choice of concealed or exposed refuge sites by preschool children viewing a model leopard in a playground simulation of antipredator behavior". International Journal of Psychological Research. 9 (2): 8–19. doi:10.21500/20112084.2325. S2CID 67792831 – via revistas.usb.edu.co.
  24. ^ Coss, Richard G. (September 1, 2021). "Something Scary Is Out There: Remembrances of Where the Threat Was Located by Preschool Children and Adults with Nighttime Fear". Evolutionary Psychological Science. 7 (3): 239–253. doi:10.1007/s40806-021-00279-9. S2CID 233619325.
  25. ^ Coss, Richard G.; Blozis, Shelley A. (December 1, 2021). "Something Scary is Out There II: the Interplay of Childhood Experiences, Relict Sexual Dinichism, and Cross-cultural Differences in Spatial Fears". Evolutionary Psychological Science. 7 (4): 359–379. doi:10.1007/s40806-021-00289-7. S2CID 237851872.
  26. ^ Coss, Richard G. (December 1, 1990). "All that Glistens: Water Connotations in Surface Finishes". Ecological Psychology. 2 (4): 367–380. doi:10.1207/s15326969eco0204_3 – via Taylor and Francis+NEJM.
  27. ^ Coss, Richard G.; Ruff, Saralyn; Simms, Tara (July 1, 2003). "All That Glistens: II. The Effects of Reflective Surface Finishes on the Mouthing Activity of Infants and Toddlers". Ecological Psychology. 15 (3): 197–213. doi:10.1207/S15326969ECO1503_1. S2CID 144073989 – via Taylor and Francis+NEJM.
  28. ^ Coss, Richard G.; Keller, Craig M. (June 1, 2022). "Transient decreases in blood pressure and heart rate with increased subjective level of relaxation while viewing water compared with adjacent ground". Journal of Environmental Psychology. 81: 101794. doi:10.1016/j.jenvp.2022.101794. S2CID 247490489.
  29. ^ Coss, Richard G. (December 1, 1991). "Context and Animal Behavior III: The Relationship Between Early Development and Evolutionary Persistence of Ground Squirrel Antisnake Behavior". Ecological Psychology. 3 (4): 277–315. doi:10.1207/s15326969eco0304_1 – via Taylor and Francis+NEJM.
  30. ^ Coss, Richard G.; Cavanaugh, Cailey; Brennan, Whitney (March 9, 2019). "Development of snake-directed antipredator behavior by wild white-faced capuchin monkeys: III. the signaling properties of alarm-call tonality". American Journal of Primatology. 81 (3): e22950. doi:10.1002/ajp.22950. PMID 30664280. S2CID 58643631.
  31. ^ Ramakrishnan, Uma; Coss, Richard G.; Schank, Jeffrey; Dharawat, Amita; Kim, Susan (April 9, 2005). "Snake Species Discrimination by Wild Bonnet Macaques (Macaca radiata)". Ethology. 111 (4): 337–356. Bibcode:2005Ethol.111..337R. doi:10.1111/j.1439-0310.2004.01063.x.
  32. ^ Ramakrishnan, Uma; Coss, Richard (January 1, 2000). "Perceptual Aspects of Leopard Recognition by Wild Bonnet Macaques (Macaca Radiata)". Behaviour. 137 (3): 315–335. doi:10.1163/156853900502105 – via brill.com.
  33. ^ Coss, Richard G.; Ramakrishnan, Uma; Schank, Jeffrey (February 28, 2005). "Recognition of partially concealed leopards by wild bonnet macaques (Macaca radiata): The role of the spotted coat". Behavioural Processes. 68 (2): 145–163. doi:10.1016/j.beproc.2004.12.004. PMID 15686825. S2CID 2256424 – via ScienceDirect.
  34. ^ Coss, Richard G.; Charles, Eric P. (March 9, 2021). "The Saliency of Snake Scales and Leopard Rosettes to Infants: Its Relevance to Graphical Patterns Portrayed in Prehistoric Art". Frontiers in Psychology. 12: 763436. doi:10.3389/fpsyg.2021.763436. PMC 8645795. PMID 34880813.
  35. ^ Coss, Richard G. (April 26, 2010). "Development of Face Aversion by the Jewel Fish (Hemichromis bimaculatus, Gill 1862)". Zeitschrift für Tierpsychologie. 48 (1): 28–46. doi:10.1111/j.1439-0310.1978.tb00246.x.
  36. ^ Coss, Richard G. (July 9, 1979). "Delayed plasticity of an instinct: Recognition and avoidance of 2 facing eyes by the jewel fish". Developmental Psychobiology. 12 (4): 335–345. doi:10.1002/dev.420120408. PMID 456760.
  37. ^ Coss, Richard G.; Globus, Albert (1978). "Spine Stems on Tectal Interneurons in Jewel Fish Are Shortened by Social Stimulation". Science. 200 (4343): 787–790. Bibcode:1978Sci...200..787C. doi:10.1126/science.644322. JSTOR 1746636. PMID 644322 – via JSTOR.
  38. ^ Coss, Richard G.; Globus, Albert (July 9, 1979). "Social experience affects the development of dendritic spines and branches on tectal interneurons in the jewel fish". Developmental Psychobiology. 12 (4): 347–358. doi:10.1002/dev.420120409. PMID 456761.
  39. ^ Wesley Burgess, J.; Coss, Richard G. (May 5, 1983). "Rapid effect of biologically relevant stimulation on tectal neurons: changes in dendritic spine morphology after nine minutes are retained for twenty-four hours". Brain Research. 266 (2): 217–223. doi:10.1016/0006-8993(83)90652-2. PMID 6871659. S2CID 24718525 – via ScienceDirect.
  40. ^ Coss, Richard G.; Perkel, Donald H. (September 1, 1985). "The function of dendritic spines: A review of theoretical issues". Behavioral and Neural Biology. 44 (2): 151–185. doi:10.1016/S0163-1047(85)90170-0. PMID 2415102 – via ScienceDirect.
  41. ^ Coss, Richard G.; Brandon, John G.; Globus, Albert (June 16, 1980). "Changes in morphology of dendritic spines on honeybee calycal interneurons associated with cumulative nursing and foraging experiences". Brain Research. 192 (1): 49–59. doi:10.1016/0006-8993(80)91007-0. PMID 7378790. S2CID 26089375 – via ScienceDirect.
  42. ^ Brandon, John G.; Coss, Richard G. (December 2, 1982). "Rapid dendritic spine stem shortening during one-trial learning: The honeybee's first orientation flight". Brain Research. 252 (1): 51–61. doi:10.1016/0006-8993(82)90977-5. PMID 7172021. S2CID 7847851 – via ScienceDirect.
  43. ^ Brubaker, Alexali S.; Coss, Richard G. (March 9, 2015). "Evolutionary constraints on equid domestication: Comparison of flight initiation distances of wild horses (Equus caballus ferus) and plains zebras (Equus quagga)". Journal of Comparative Psychology. 129 (4): 366–376. doi:10.1037/a0039677. PMID 26348970.
  44. ^ Brubaker, Alexali S.; Coss, Richard G. (July 9, 2016). Ebensperger, L. (ed.). "Effects of Single- and Mixed-Species Group Composition on the Flight Initiation Distances of Plains and Grevy's Zebras". Ethology. 122 (7): 531–541. doi:10.1111/eth.12500. S2CID 87346253.
  45. ^ Coss, Richard G. (2017). "Drawings of Representational Images by Upper Paleolithic Humans and their Absence in Neanderthals Reflect Historical Differences in Hunting Wary Game". Evolutionary Studies in Imaginative Culture. 1 (2): 15–38. doi:10.26613/esic.1.2.46. JSTOR 10.26613/esic.1.2.46. S2CID 134391188.
  46. ^ "Effects of predator behavior and proximity on risk assessment by Columbian black-tailed deer".
  47. ^ "Effects of risk assessment, predator behavior, and habitat on escape behavior in Columbian black-tailed deer".
  48. ^ Ramakrishnan, Uma; Coss, Richard G. (July 1, 2001). "Strategies used by bonnet macaques (Macaca radiata) to reduce predation risk while sleeping". Primates. 42 (3): 193–206. doi:10.1007/BF02629636. S2CID 34596783 – via Springer Link.
  49. ^ "A Comparison of the Sleeping Behavior of Three Sympatric Primates".
  50. ^ Ramakrishnan, Uma; Coss, Richard G.; Pelkey, Neil W. (July 1, 1999). "Tiger decline caused by the reduction of large ungulate prey: evidence from a study of leopard diets in southern India". Biological Conservation. 89 (2): 113–120. Bibcode:1999BCons..89..113R. doi:10.1016/S0006-3207(98)00159-1 – via ScienceDirect.
  51. ^ Thuppil, Vivek; Coss, Richard G. (April 9, 2016). "Playback of felid growls mitigates crop-raiding by elephants Elephas maximus in southern India". Oryx. 50 (2): 329–335. doi:10.1017/S0030605314000635. S2CID 86369635.
  52. ^ Thuppil, Vivek; Coss, Richard G. (April 1, 2012). "Using Threatening Sounds as a Conservation Tool: Evolutionary Bases for Managing Human–Elephant Conflict in India". Journal of International Wildlife Law & Policy. 15 (2): 167–185. doi:10.1080/13880292.2012.678794. S2CID 85122785 – via Taylor and Francis+NEJM.
  53. ^ Thuppil, Vivek; Coss, Richard G. (October 23, 2013). "Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger". Biology Letters. 9 (5): 20130518. doi:10.1098/rsbl.2013.0518. PMC 3971691. PMID 24026347.
  54. ^ Coss, R. G.; Burgess, J. W. (September 28, 1981). "Jewel fish retain juvenile schooling pattern after crowded development". Developmental Psychobiology. 14 (5): 451–457. doi:10.1002/dev.420140507. PMID 7196851 – via PubMed.
  55. ^ Burgess, J. W.; Coss, R. G. (July 28, 1981). "Short-term juvenile crowding arrests the developmental formation of dendritic spines on tectal interneurons in jewel fish". Developmental Psychobiology. 14 (4): 389–396. doi:10.1002/dev.420140412. PMID 7250527 – via PubMed.
  56. ^ Burgess, J. W.; Coss, R. G. (1982). "Effects of chronic crowding stress on midbrain development: Changes in dendritic spine density and morphology in jewel fish optic tectum". Developmental Psychobiology. 15 (5): 461–470. doi:10.1002/dev.420150508. PMID 6890000.