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Encephalization quotient

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Species Encephalization quotient (EQ)[1]
Bottlenose dolphin4.14[2]
Rhesus monkey2.1

Encephalization Quotient (EQ), or encephalization level is a measure of relative brain size defined as the ratio between actual brain mass and predicted brain mass for an animal of a given size, which is hypothesized to be a rough estimate of the intelligence of the animal.[5]

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.[6]

Brain-body size relationship

Brain size usually increases with body size in animals (is positively correlated), i.e. large animals usually have larger brains than smaller animals.[7] 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 comparatively small brain/body size (1/560), despite elephants being obviously quite intelligent animals.[7][8]

There are possibly several reasons for this trend, but one of them is that neural cells have a relative constant size. As an animal's brain gets larger, addition of more nerve cells will cause the brain to increase in size to a lesser degree than the rest of the body. This phenomenon has been called the cephalization factor; E = CS2, where E and S are body and brain weights and C is the cephalization factor.[9] Thus just focusing on the relationship between the body and the brain is not enough; one also has to consider the total size of the animal. To compensate for this factor, a formula has been devised by plotting the brain/body weight of various mammals against each other and a curve fitted so as to give best fit to the data.

The formula for the curve varies, but is usually given as Ew(brain) = 0.12w(body)2/3.[6] 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.[6]

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. It also puts humans at the top of the list. The relationship between brain-to-body mass ratio and complexity of behaviour is not perfect as other factors also influence intelligence, like the evolution of the recent cerebral cortex and different degrees of brain folding,[10] which increase the surface of the cortex, which is positively correlated to intelligence in humans.[11]

Comparisons between groups

Dolphins have the highest brain-to-body weight ratio of all cetaceans.[3] Manta rays have the highest for a fish[12], and either octopuses[9] or jumping spiders[13] have the highest for an invertebrate. 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.[14][15][16] Mean EQ for reptiles are about one tenth of the EQ for 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.[17] Estimation of brain size in the oldest known bird, Archaeopteryx, shows it had an EQ in the upper reptilian range, but below that of living birds.[18]

Species Simple brain-to body ratio (E/S)[7]
small birds1/12

In the essay "Bligh's Bounty",[19] Stephen Jay Gould 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. This formula is useless for invertebrates because they do not have spinal cords, or in some cases, central nervous systems.


Recent research indicates that whole brain size is a better measure of cognitive abilities than EQ for primates at least.[20]

The concept of EQ as a measure of intelligence can be strongly criticised in a very simple argument. The brains of large dinosaurs were frequently tiny.[21] Stegosaurus, weighing about the same as an average elephant, had a comparatively small brain—160 g compared to about 5 kg for an elephant. While Stegosaurus undoubtedly was an animal of very limited behavioural complexity, this fact undermines the idea on which EQ is based - that a larger animal requires a larger brain to look after a large body.

If Stegosaurus could survive with this tiny brain, it would seem that any animal with anything bigger must be using it for non-essential abilities. However, mammalian evolution has repeatedly improved the effectiveness of a bodily function by innervating it more; the digestive and immune systems are examples. Thus, while an elephant has a much larger brain than a Stegosaurus, a substantial part of the excess brain is bound up in bodily functions rather than cognitive functions. This can account for some differences between classes of animals, but not species within a class. More associative brain tissue, cortex, still indicates a level of mental activity above the reptilian form. Some of these abilities may be sensory and/or physical, and some may be intellectual. The actual intelligence of an animal therefore depends on the size of the brain and the proportion of the brain that is used for intellectual abilities, rather than advanced sensory or physical skills. Critics[attribution needed] point out that EQ gives only a very rough estimate of these proportions.

See also


  1. Gerhard Roth und Ursula Dicke (2005-05). Evolution of the brain and Intelligence. TRENDS in Cognitive Sciences 9 (5): 250.
  2. 2.0 2.1 Marino, Lori (2004). Cetacean Brain Evolution: Multiplication Generates Complexity. International Society for Comparative Psychology (17): 1–16.
  3. 3.0 3.1 Marino, L. and Sol, D. and Toren, K. and Lefebvre, L. (2006). Does diving limit brain size in cetaceans?. Marine Mammal Science 22 (2): 413–425.
  4. Shoshani, Jeheskel (30 June). Elephant brain Part I: Gross morphology, functions,comparative anatomy, and evolution. Brain Research Bulletin 70 (2): 124–157.
  5. G.Rieke. Natural Sciences 102: Lecture Notes: Emergence of Intelligence. URL accessed on 2011-02-12.
  6. 6.0 6.1 6.2 Moore, J. (1999): Allometry, University of California, San Diego
  7. 7.0 7.1 7.2
  8. Hart, B.L., L.A. Hart, M. McCoy, C.R. Sarath (November 2001). Cognitive behaviour in Asian elephants: use and modification of branches for fly switching. Animal Behaviour 62 (5): 839–847.
  9. 9.0 9.1 Gould (1977)Ever since Darwin, c7s1
  10. Cortical Folding and Intelligence. URL accessed on 2008-09-15.
  11. Haier, R.J., Jung, R.E., Yeo, R.C., Head, K. and Alkired, M.T. (2004): Structural brain variation and general intelligence. NeuroImage Vol. 23, Issue 1, September 2004, Pages 425-433 summary
  12. Striedter, Georg F. (2005). Principles of brain evolution, Sunderland, Mass.: Sinauer.
  13. Jumping Spider Vision. URL accessed on 2009-10-28.
  14. 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.
  15. James K. Riling (1999). The Primate Neocortex in Comparative Perspective using Magnetic Resonance Imaging. Journal of Human Evolution 37 (2): 191–223.
  16. Suzana Herculano-Houzel (2009). The Human Brain in Numbers- A Linearly Scaled-Up Primae Brain. Frontiers in Human Neuroscience 2: 1–11 (2).
  17. Paul, Gregory S. (1988) Predatory dinosaurs of the world. Simon and Schuster. ISBN 0671619462
  18. Hopson J.A. (1977). Relative Brain Size and Behavior in Archosaurian Reptiles. Annual Review of Ecology and Systematics 8: 429–448.
  19. web archive of
  20. Overall Brain Size, and Not Encephalization Quotient, Best Predicts Cognitive Ability across Non-Human Primates. Brain Behav Evol 2007;70:115-124 (DOI: 10.1159/000102973)[1]
  21. [2]

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