Encephalization quotient

Species Encephalization
quotient (EQ)[1]
Human7.4–7.8
Tucuxi4.56[2]
Bottlenose dolphin4.14[3]
Orca2.57–3.3[3][4]
Chimpanzee2.2–2.5[5]
Rhesus macaque2.1
Elephant1.13–2.36[6]
Chinchilla 1.34[7]
Dog1.2
Squirrel1.1
Cat1.00
Horse0.9
Sheep0.8
Taurine cattle0.52-0.59[8]
Mouse0.5
Rat0.4
Rabbit0.4

Encephalization quotient (EQ), encephalization level or just encephalization is a relative brain size measure that is defined as the ratio between actual brain mass and predicted brain mass for an animal of a given size, which may approximate intelligence level or cognition of the species.[9][10]

This is a more refined measurement than the raw brain-to-body mass ratio, as it takes into account allometric effects. The relationship, expressed as a formula, has been developed for mammals, and may not yield relevant results when applied outside this group.[11] Additionally to volume, mass or cell count, the energy expenditure of the brain could be compared with that of the rest of the body.

Encephalization can also refer to the tendency for a species to evolve larger brains through time. Anthropological studies indicate that bipedalism preceded encephalization in the human evolutionary lineage after divergence from the chimpanzee lineage. Compared to the chimpanzee brain, the human brain is larger and certain brain regions have been particularly altered during human evolution.[12] Most brain growth of chimpanzees happens before birth while most human brain growth happens after birth.[13] In 2004, Dennis Bramble and Daniel Lieberman proposed that early Homo were scavengers who used stone tools to harvest meat off carcasses and to open bones. They proposed that humans specialized in long-distance running to compete with other scavengers in reaching carcasses.[14] It has been suggested that such an adaptation ensured a food supply that made large brains possible.[15] More encephalized species tend to have longer spinal shock duration.


Brain-body size relationship

Species Simple brain-to-body
ratio (E/S)[16]
small birds1/12
human1/40
mouse1/40
dolphin1/50
cat1/100
chimpanzee1/113
dog1/125
frog1/172
lion1/550
elephant1/560
horse1/600
shark1/2496
hippopotamus1/2789

Brain size usually increases with body size in animals (is positively correlated), i.e. large animals usually have larger brains than smaller animals.[16] The relationship is not linear, however. Generally, small mammals have relatively larger brains than big ones. Mice have a direct brain/body size ratio similar to humans (1/40), while elephants have a comparatively small brain/body size (1/560), despite being quite intelligent animals.[16][17]

Several reasons for this trend are possible, one of which is that neural cells have a relative constant size.[18] Some brain functions, like the brain pathway responsible for a basic task like drawing breath, are basically similar in a mouse and an elephant. Thus, the same amount of brain matter can govern breathing in a large or a small body. While not all control functions are independent of body size, some are, and hence large animals need comparatively less brain than small animals. This phenomenon has been called the cephalization factor: C = E ÷ S2 , where E and S are brain and body weights respectively, and C is the cephalization factor.[19] To determine the formula for this factor, the brain/body weights of various mammals were plotted against each other, and the curve of E = C × S2 chosen as the best fit to that data.[20]

The cephalization factor and the subsequent encephalization quotient was developed by H.J. Jerison in the late 1960s.[21] The formula for the curve varies, but an empirical fitting of the formula to a sample of mammals gives .[11] As this formula is based on data from mammals, it should be applied to other animals with caution. For some of the other vertebrate classes the power of 3/4 rather than 2/3 is sometimes used, and for many groups of invertebrates the formula may give no meaningful results at all.[11]

Calculation

Snell's equation of simple allometry[22] is:

Here "E" is the weight of the brain, "C" is the cephalization factor and "S" is body weight and "r" is the exponential constant. The exponential constant for primates is 0.28[22] and either 0.56 or 0.66 for mammals in general.[23]

The "Encephalization Quotient" (EQ) is the coefficient "C" in Snell's allometry equation, usually normalized with respect to a reference species. In the following table, the coefficients have been normalized with respect to the value for the cat, which is therefore attributed an EQ of 1 .[23]

Another way to calculate encephalization quotient is by dividing the actual weight of an animal's brain with its predicted weight according to Jerison's formula.

Species EQ[23] Species EQ[23]
Human 7.44 Dog 1.17
Dolphin 5.31 Cat 1.00
Chimpanzee 2.49 Horse 0.86
Raven[24] 2.49 Sheep 0.81
Rhesus monkey 2.09 Mouse 0.50
Elephant 1.87 Rat 0.40
Baleen whale 1.76 Rabbit 0.40

This measurement of approximate intelligence is more accurate for mammals than for other classes and phyla of Animalia.

EQ and intelligence in mammals

Intelligence in animals is hard to establish, but the larger the brain is relative to the body, the more brain weight might be available for more complex cognitive tasks. The EQ formula, as opposed to the method of simply measuring raw brain weight or brain weight to body weight, makes for a ranking of animals that coincide better with observed complexity of behaviour.

Mean EQ for mammals is around 1, with carnivorans, cetaceans and primates above 1, and insectivores and herbivores below. This reflects two major trends. One is that brain matter is extremely costly in terms of energy needed to sustain it.[25] Animals which live on relatively nutrient poor diets (plants, insects) have relatively little energy to spare for a large brain, while animals living from energy-rich food (meat, fish, fruit) can grow larger brains. The other factor is the brain power needed to catch food. Carnivores generally need to find and kill their prey, which presumably requires more cognitive power than browsing or grazing.[26][27] The brain size of a wolf is about 30% larger than a similarly sized domestic dog, again reflecting different needs in their respective way of life.[28]

It is worth noting, however, that of the animals demonstrating the highest EQ's (see associated table), many are primarily herbivorous, including apes, macaques, and proboscideans. The dietary factor, therefore, may be less significant than certain others, like gregariousness.

Another factor affecting relative brain size is sociality and flock size.[29] For example, dogs (a social species) have a higher EQ than cats (a mostly solitary species). Animals with very large flock size and/or complex social systems consistently score high EQ, with dolphins and orcas having the highest EQ of all cetaceans,[4] and humans with their extremely large societies and complex social life topping the list by a good margin.[1]

Comparisons with non-mammalian animals

Birds generally have lower EQ than mammals, but parrots and particularly the corvids show remarkable complex behaviour and high learning ability. Their brains are at the high end of the bird spectrum, but low compared to mammals. Bird cell size is on the other hand generally smaller than that of mammals, which may mean more brain cells and hence synapses per volume, allowing for more complex behaviour from a smaller brain.[1] Both bird intelligence and brain anatomy are however very different from those of mammals, making direct comparison difficult.[30]

Manta rays have the highest EQ among fish,[31] and either octopuses[19] or jumping spiders[32] have the highest among invertebrates. Despite the jumping spider having a huge brain for its size, it is minuscule in absolute terms, and humans have a much higher EQ despite having a lower raw brain-to-body weight ratio.[33][34][35] Mean EQs for reptiles are about one tenth of those of mammals. EQ in birds (and estimated EQ in dinosaurs) generally also falls below that of mammals, possibly due to lower thermoregulation and/or motor control demands.[36] Estimation of brain size in the oldest known bird, Archaeopteryx, shows it had an EQ well above the reptilian range, and just below that of living birds.[37]

Biologist Stephen Jay Gould has noted that if one looks at vertebrates with very low encephalization quotients, their brains are slightly less massive than their spinal cords. Theoretically, intelligence might correlate with the absolute amount of brain an animal has after subtracting the weight of the spinal cord from the brain.[38] This formula is useless for invertebrates because they do not have spinal cords or, in some cases, central nervous systems.

EQ in paleoneurology

Behavioural complexity in living animals can to some degree be observed directly, making the predictive power of the encephalization quotient less relevant. It is however central in paleoneurology, where the endocast of the brain cavity and estimated body weight of an animal is all one has to work from. The behaviour of extinct mammals and dinosaurs is typically investigated using EQ formulas.[21]

Encephalization quotient is also used in estimating evolution of intelligent (human) behaviour in human ancestors. Fossils of archaic humans have EQ e.g. 4.15.[39]

Criticism

Recent research indicates that whole brain size is a better measure of cognitive abilities than EQ for non-human primates at least.[40] The relationship between brain-to-body mass ratio and complexity is not alone in influencing intelligence. Other factors, such as the recent evolution of the cerebral cortex and different degrees of brain folding,[41] which increases the surface area (and volume) of the cortex, are positively correlated to intelligence in humans.[42]

History

Quantifying an animal's encephalization has been argued to be directly proportional, although not equal, to that animal's level of intelligence. Aristotle wrote in 335 BCE: "Of all the animals, man has the brain largest in proportion to his size."[43] Also, in 1871, Charles Darwin wrote in his book The Descent of Man: "No one, I presume, doubts that the large proportion which the size of man's brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his mental powers."[44]

See also

References

  1. 1 2 3 Gerhard Roth und Ursula Dicke (May 2005). "Evolution of the brain and Intelligence". Trends in Cognitive Sciences. 9 (5): 250–7. doi:10.1016/j.tics.2005.03.005. PMID 15866152.
  2. ↑ William F. Perrin; Bernd Würsig; J.G.M. Thewissen (2009). Encyclopedia of Marine Mammals. Academic Press. p. 150. ISBN 978-0-08-091993-5.
  3. 1 2 Marino, Lori (2004). "Cetacean Brain Evolution: Multiplication Generates Complexity" (PDF). International Society for Comparative Psychology (17): 1–16. Retrieved 2010-08-29.
  4. 1 2 Marino, L. and Sol, D. and Toren, K. and Lefebvre, L. (2006). "Does diving limit brain size in cetaceans?" (PDF). Marine Mammal Science. 22 (2): 413–425. doi:10.1111/j.1748-7692.2006.00042.x.
  5. ↑ Hill, Kyle. "How science could make a chimp like Dawn of the Planet of the Apes' Caesar's". The Nerdist. Retrieved 10 December 2014.
  6. ↑ Shoshani, Jeheskel; Kupsky, William J.; Marchant, Gary H. (30 June 2006). "Elephant brain Part I: Gross morphology, functions, comparative anatomy, and evolution". Brain Research Bulletin. 70 (2): 124–157. doi:10.1016/j.brainresbull.2006.03.016. PMID 16782503.
  7. ↑ Spotorno, Angel E.; Zuleta, Carlos A.; Valladares, J. Pablo; Deane, Amy L.; Jiménez, Jaime E. (2004-12-01). "Chinchilla laniger". Mammalian Species. 758 (758): 1–9. doi:10.2307/3504402 (inactive 2018-09-23). JSTOR 3504402.
  8. ↑ Raia, Pasquale; Ballarin, Cristina; Povinelli, Michele; Granato, Alberto; Panin, Mattia; Corain, Livio; Peruffo, Antonella; Cozzi, Bruno (2016). "The Brain of the Domestic Bos taurus: Weight, Encephalization and Cerebellar Quotients, and Comparison with Other Domestic and Wild Cetartiodactyla". PLOS One. 11 (4): e0154580. Bibcode:2016PLoSO..1154580B. doi:10.1371/journal.pone.0154580. ISSN 1932-6203. PMC 4851379. PMID 27128674.
  9. ↑ Pontarotti, Pierre (2016). Evolutionary Biology: Convergent Evolution, Evolution of Complex Traits. Springer. p. 74. ISBN 978-3-319-41324-2.
  10. ↑ G.Rieke. "Natural Sciences 102: Lecture Notes: Emergence of Intelligence". Retrieved 2011-02-12.
  11. 1 2 3 Moore, J. (1999): Allometry, University of California, San Diego
  12. ↑ See Figures 1 and 2 of Bradbury J (March 2005), "Molecular insights into human brain evolution", PLoS Biol., 3 (3): e50, doi:10.1371/journal.pbio.0030050, PMC 1065704, PMID 15760271.
  13. ↑ Penin, X; Berge, C; Baylac, M (2002). "Ontogenetic study of the skull in modern humans and the common chimpanzees: Neotenic hypothesis reconsidered with a tridimensional Procrustes analysis". American Journal of Physical Anthropology. 118 (1): 50–62. doi:10.1002/ajpa.10044. PMID 11953945.
  14. ↑ Bramble DM, Lieberman DE (November 2004), "Endurance running and the evolution of Homo" (PDF), Nature, 432 (7015): 345–52, Bibcode:2004Natur.432..345B, doi:10.1038/nature03052, PMID 15549097.
  15. ↑ Aiello LC, Wheeler P (April 1995), "The Expensive-Tissue Hypothesis: The Brain and the Digestive System in Human and Primate Evolution" (PDF), Current Anthropology, 36 (2): 199–221, doi:10.1086/204350.
  16. 1 2 3 http://serendip.brynmawr.edu/bb/kinser/Int3.html
  17. ↑ Hart, B. L.; Hart, L. A.; McCoy, M.; Sarath, C. R. (November 2001). "Cognitive behaviour in Asian elephants: use and modification of branches for fly switching". Animal Behaviour. 62 (5): 839–847. doi:10.1006/anbe.2001.1815. Retrieved 2007-10-30.
  18. ↑ Oxnard, C.; Cartmill, M.; Brown, K.B. (2008). The human body : developmental, functional and evolutionary bases. Hoboken, N.J.: Wiley. p. 274. ISBN 978-0471235996.
  19. 1 2 Gould (1977) Ever since Darwin, c7s1
  20. ↑ Jerison, H.J. (1983). Eisenberg, J.F.; Kleiman, D.G., eds. Advances in the Study of Mammalian Behavior. Pittsburgh: Special Publication of the American Society of Mammalogists, nr. 7. pp. 113–146.
  21. 1 2 Brett-Surman, Michael K.; Holtz, Thomas R.; Farlow, James O., eds. (2012-06-27). The complete dinosaur. Illustrated by Bob Walters (2nd ed.). Bloomington, Ind.: Indiana University Press. pp. 191–208. ISBN 978-0-253-00849-7.
  22. 1 2 Williams, M.F. (April 2002), "Primate encephalization and intelligence", Medical Hypotheses, 58 (4): 284–290, doi:10.1054/mehy.2001.1516, PMID 12027521
  23. 1 2 3 4 "Thinking about brain size". Serendip Studeio. Retrieved 21 May 2011.
  24. ↑ Emery, N. J. (2006). "Cognitive ornithology: The evolution of avian intelligence". Philosophical Transactions of the Royal Society B: Biological Sciences. 361 (1465): 23–43. doi:10.1098/rstb.2005.1736. PMC 1626540. PMID 16553307.
  25. ↑ Isler, K.; van Schaik; C. P (22 December 2006). "Metabolic costs of brain size evolution". Biology Letters. 2 (4): 557–560. doi:10.1098/rsbl.2006.0538. PMC 1834002. PMID 17148287.
  26. ↑ Savage, J.G. (1977). "Evolution in carnivorous mammals" (PDF). Palaentology. 20, part 2: 237–271. Archived from the original (PDF) on 13 December 2013. Retrieved 19 February 2013.
  27. ↑ Lefebvre, Louis; Reader, Simon M.; Sol, Daniel (1 January 2004). "Brains, Innovations and Evolution in Birds and Primates". Brain, Behavior and Evolution. 63 (4): 233–246. doi:10.1159/000076784. PMID 15084816. Retrieved 19 February 2013.
  28. ↑ Horowitz, A. "Why Brain Size Doesn't Correlate With Intelligence". Smithsonian.com. Retrieved 29 January 2016.
  29. ↑ Susanne Shultz; R.I.M. Dunbar (2006). "Both social and ecological factors predict ungulate brain size". Proceedings of the Royal Society B: Biological Sciences. 273 (1583): 207–215. doi:10.1098/rspb.2005.3283. PMC 1560022. PMID 16555789.
  30. ↑ Emery, N. J. (29 January 2006). "Cognitive ornithology: the evolution of avian intelligence". Philosophical Transactions of the Royal Society B: Biological Sciences. 361 (1465): 23–43. doi:10.1098/rstb.2005.1736. PMC 1626540. PMID 16553307.
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  33. ↑ Meyer, W., Schlesinger, C., Poehling, H.M. & Ruge, W. (1984): Comparative and quantitative aspects of putative neurotransmitters in the central nervous system of spiders (Arachnida: Araneida). Comparative Biochemical Physiology no 78 (C series): pp 357–62.
  34. ↑ James K. Riling; Insel, TR (1999). "The Primate Neocortex in Comparative Perspective using Magnetic Resonance Imaging". Journal of Human Evolution. 37 (2): 191–223. doi:10.1006/jhev.1999.0313. PMID 10444351.
  35. ↑ Suzana Herculano-Houzel (2009). "The Human Brain in Numbers- A Linearly Scaled-Up Primate Brain". Frontiers in Human Neuroscience. 3: 1–11 (2). doi:10.3389/neuro.09.031.2009. PMC 2776484. PMID 19915731.
  36. ↑ Paul, Gregory S. (1988) Predatory dinosaurs of the world. Simon and Schuster. ISBN 0-671-61946-2
  37. ↑ Hopson J.A. (1977). "Relative Brain Size and Behavior in Archosaurian Reptiles". Annual Review of Ecology and Systematics. 8: 429–448. doi:10.1146/annurev.es.08.110177.002241.
  38. ↑ "Bligh's Bounty". Archived from the original on 9 July 2001. Retrieved 12 May 2011.
  39. ↑ http://www.pnas.org/content/103/10/3552.full - Body size, body proportions, and encephalization in a Middle Pleistocene archaic human from northern China
  40. ↑ Deaner, Robert O; Isler, Karin; Burkart, Judith; Van Schaik, Carel (2007). "Overall Brain Size, and Not Encephalization Quotient, Best Predicts Cognitive Ability across Non-Human Primates". Brain Behav Evol. 70 (2): 115–124. CiteSeerX 10.1.1.570.7146. doi:10.1159/000102973. PMID 17510549.
  41. ↑ "Cortical Folding and Intelligence". Retrieved 2008-09-15.
  42. ↑ Haier, R.J., Jung, R.E., Yeo, R.C., Head, K. and Alkired, M.T. (Sep 2004). "Structural brain variation and general intelligence". NeuroImage. 23 (1): 425–33. doi:10.1016/j.neuroimage.2004.04.025. PMID 15325390.
  43. ↑ Russell, Stuart; Norvig, Peter (2003), Artificial Intelligence: A Modern Approach, Upper Saddle River, N.J.: Prentice Hall/Pearson Education, ISBN 978-0-13-790395-5
  44. ↑ Darwin, Charles (1981), The Descent of Man, and Selection in Relation to Sex (1981 reprint of 1871 ed.), Princeton, New Jersey: Princeton University Press, p. 145, ISBN 978-0-691-02369-4 See also quote, p.60, in online text of earlier reprint of second (1874) edition.

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