Given the criteria we have for considering whether a being is conscious, it is reasonable to conclude that vertebrates and a large number of invertebrates are conscious. The clearer cases are those of animals who have a centralized nervous system whose central organ (basically, a brain) has some development. However, there are a number of animals who possess centralized nervous systems whose central organ is not quite developed. In these cases, doubts may arise about whether they are conscious or not. The reason is, if in order to be conscious, it is necessary that a nervous system be organized a certain way, then the evolutionary path that will lead there will necessarily pass first, in its previous stages, through the existence of a nervous system without any centralization, and afterwards through a nervous system that starts to be centralized, but not enough to support consciousness. First, the nervous system becomes minimally centralized, with some very simple nervous ganglia, then, with more complex ganglia. Nervous systems become more complex until, at some point, the phenomenon of consciousness appears. Along the evolutionary path, there may be stages where there are some minimally centralized nervous systems that don’t give rise to consciousness.
We don’t know with full certainty if there are currently animals with minimally centralized nervous systems that don’t give rise to consciousness. It may be that all the centralized nervous systems that exist currently are centralized enough to host consciousness. This would be the case if all those that were in an intermediate stage, that is, having minimally centralized nervous systems that don’t give rise to consciousness, were already extinct. We have no answer to this question at this point.
Among animals that are conscious, we can count with a high degree of certainty vertebrates including human beings and invertebrates such as cephalopods (such as octopuses and squids), since they satisfy the criteria for sentience. In addition, we also have strong reasons to think that other animals such as arthropods (insects, arachnids, and crustaceans) are conscious too. The physiology of these animals is organized in ways that seem to be sufficient for giving rise to consciousness, and their behavior also seems to support this.1
As for other animals, such as bivalve mollusks, we don’t have reasons as strong as those we have in the previous cases.2 However, given the problems involved in determining the basis of consciousness, we cannot rule out completely the possibility that they are sentient.
The following are some examples of animals who would fall, respectively, in these two groups.
It is a controversial issue whether animals such as insects, arachnids and other arthropods are sentient.3
In the case of insects, we can consider the following line of reasoning, which is an argument by homology. Insects possess a centralized nervous system that is centralized not merely due to the presence of ganglia, but actually includes a brain. It must be noted, though, that it is a very simple and small brain. Therefore, considering insects’ physiology alone is not enough to conclude whether they are conscious or not. Apart from this, the behavior of some insects is very simple. Others, however, have very complex behavior. A clear example of this is bees. Their behavior, including their famous waggle dance, leads us to think that they really are beings with experiences, that is, they are conscious.4 There are other insects that have a very similar physiological structure to that of bees but that exhibit only much simpler behaviors, such as mosquitoes. Because of the similarity of their nervous systems, we might believe that if bees are conscious, then they are conscious, too. We must bear in mind, though, that this does not follow automatically. We must not lose sight of the fact that insects are the most numerous class of animals currently existing. Due to this, there are certain differences among them that are much more significant than those that occur among mammals, for instance.
Because of this greater variation among insects, a different response may be to claim that bees (or, in general, hymenopterans, the insect order to which bees belong and which includes wasps and ants) are conscious, while other insects are not. Or, maybe, that even if all insects are conscious, bees are able to have more vivid experiences. This seems more likely to be the case than that only some insects are sentient. Although the differences in the behaviors of insects are significant, the differences between their physiologies are not so important as to lead us to conclude that only some of them are sentient.
Of course, a different line of reasoning is also possible. We might think that beings exhibiting only simple behaviors could not be sentient. From here, we could posit that the structure of the nervous systems of these animals would not be complex enough for consciousness to appear (despite its centralization). Therefore, we would conclude that, since their nervous systems are similar to those of animals exhibiting only simple behaviors, animals such as bees would not really be conscious, since they would lack the necessary nervous structure. We would then claim that even behaviors as complex as those of bees could occur through mechanisms that would not imply the presence of consciousness. This explanation, however, seems less plausible than the previous one (that the nervous systems of all insects are similar enough that if some insects are conscious, they must all be). A being may be conscious and display a relatively simple behavior. It seems more unlikely, though, that a nonconscious being would display a complex behavior.5
In the same vein, we could consider other criteria such as the presence of natural opiates among insects. This would reinforce the claim that these animals are sentient.
In the case of other arthropods, such as arachnids, we cannot appeal to evolutionary logic to apply the conclusions we reach in the case of insects, given that they are not closely related. Despite this, we may follow an argument from homology. Insects’ nervous structures are not significantly more complex than arachnids’. Plus, the behavior of arachnids is not very different from that of a number of insects. Therefore, it may make sense to infer that if insects are sentient, then arachnids are sentient too.
We can see that we are facing a question to which we cannot arrive at an immediate and clear answer. However, we can consider all the different criteria we have to examine the question together, and ponder all the evidence we have in order to make progress towards finding which is the most plausible answer. This reasoning process is similar to the one that is followed in the case of other animals (such as vertebrates). It is only that here we may need to consider more factors.
The problem becomes more complex if we consider other beings with a simpler structure, without a brain, but only some central nervous ganglia. This is the structure of many invertebrates, such as, for example, bivalve mollusks (including mussels and oysters) and gastropods (including snails).6 The appeal to evolutionary logic in these cases is not useful, since the behavior these animals display is very simple. It could be performed without requiring that the animals that display it be conscious. This happens in particular in animals that stay attached to rocks or other surfaces without moving, in the case of bivalves or of certain crustacea such as barnacles. Bivalves can perform some movements, such as opening and closing their shells. But these movements may be triggered in a more economic way in terms of energy by some stimulus-response mechanism (in fact, their behavior is not more complex than that of other beings without a centralized nervous system, such as carnivorous plants or certain echinoderms). At any rate, their physiology leaves the question open.7 It might be that they have experiences. It is not possible to rule out that possibility given our lack of knowledge regarding how to answer the question of what is the basis of consciousness.
There are other indicators that are not conclusive, although they may help us to appraise the question. Bivalves have mechanisms that are analogous to opiate receptors possessed by other animals.8 In other animals, the function of these receptors is to make it possible to have their suffering relieved when they are in significant pain. Due to this, a very plausible explanation of why bivalves have them, maybe the most plausible one, is that they can suffer too. But this is not totally conclusive. It is also possible that the organisms of these animals use these substances with an aim that is different from the one that they have in other animals.
Apart from these, there are other reasons that support the idea that bivalves and other animals with very simple centralized nervous systems can suffer. One of them is that some bivalves have simple eyes, and the most plausible explanation is that a being with eyes also has the experience of vision (such as snails).9 It has been discovered that the heart rate of bivalves speeds up in situations in which they are threatened by predators.10 In addition, sounds and vibrations are in the sensitivity range of mussels and oysters.11 These indicators, again, are not conclusive, but they show that it is not clear that these animals are not conscious. In the case of other animals that can have nervous systems with some centralization, we can say something similar.
Allen, C. & Trestman, M. (2004) “Animal consciousness”, in Zalta, E. N. (ed.) Stanford encyclopedia of philosophy, Stanford: The Metaphysics Research Lab [accessed on 18 February 2015].
Barr, S.; Laming, P. R.; Dick, J. T. A. & Elwood, R. W. (2008) “Nociception or pain in a decapod crustacean?”, Animal Behaviour, 75, pp. 745-751.
Broom, D. M. (2007) “Cognitive ability and sentience: Which aquatic animals should be protected?”, Diseases of Aquatic Organisms, 75, pp. 99-108.
Crook, R. J. (2013) “The welfare of invertebrate animals in research: Can science’s next generation improve their lot?”, Journal of Postdoctoral Research, 1 (2), pp. 9-18 [accessed on 22 February 2014].
Crook, R. J.; Hanlon, R. T. & Walters, E. T. (2013) “Squid have nociceptors that display widespread long-term sensitization and spontaneous activity after bodily injury”, The Journal of Neuroscience, 33, pp. 10021-10026.
Dawkins, M. S. (2001) “Who needs consciousness?”, Animal Welfare, 10, pp. 19-29.
Eisemann, C. H.; Jorgensen, W. K.; Merritt, D. J.; Rice, M. J.; Cribb, B. W.; Webb, P. D. & Zalucki, M. P. (1984) “Do insects feel pain? A biological view”, Experentia, 40, pp. 164-167.
Elwood, R. W. (2011) “Pain and suffering in invertebrates?”, ILAR Journal, 52, pp. 175-184.
Elwood, R. W. & Adams, L. (2015) “Electric shock causes physiological stress responses in shore crabs, consistent with prediction of pain”, Biology Letters, 11 (1) [accessed on 13 November 2015].
Elwood, R. W. & Appel, M. (2009) “Pain experience in hermit crabs?”, Animal Behaviour, 77, pp. 1243-1246.
Fiorito, G. (1986) “Is there ‘pain’ in invertebrates?”, Behavioural Processes, 12, pp. 383-388.
Gherardi, F. (2009) “Behavioural indicators of pain in crustacean decapods”, Annali dell´Istituto Superiore di Sanita, 45, pp. 432-438.
Gentle, M. J. (1992) “Pain in birds”, Animal Welfare, 1, pp. 235-247.
Griffin, D. R. (1984) Animal thinking, Cambridge: Harvard University Press.
Griffin, D. R. (2001) Animal minds: Beyond cognition to consciousness, Chicago: Chicago University Press.
Harvey-Clark, C. (2011) “IACUC challenges in invertebrate research”, ILAR Journal, 52, pp. 213-220 [accessed on 14 February 2013].
Horvath, K.; Angeletti, D.; Nascetti, G. & Carere, C. (2013) “Invertebrate welfare: An overlooked issue”, Annali dell´Istituto superiore di sanità, 49, pp. 9-17 [accessed on 3 October 2013].
Huffard, C. L. (2013) “Cephalopod neurobiology: An introduction for biologists working in other model systems”, Invertebrate Neuroscience, 13, pp. 11-18.
Knutsson, S. (2015a) The moral importance of small animals, Master’s thesis in practical philosophy, Gothenburg: University of Gothenburg [accessed on 4 January 2016].
Knutsson, S. (2015b) “How good or bad is the life of an insect”, simonknutsson.com [accessed on 4 January 2016].
Leonard, G. H.; Bertness, M. D. & Yund, P. O. (1999) “Crab predation, waterborne cues, and inducible defenses in the blue mussel, Mytilus edulis”, Ecology, 75, pp. 1-14.
Lozada, M.; Romano, A. & Maldonado, H. (1988) “Effect of morphine and naloxone on a defensive response of the crab Chasmagnathus granulatus”, Pharmacology, Biochemistry, and Behavior, 30, pp. 635-640.
Magee, B.; Elwood, R. W. (2013) “Shock avoidance by discrimination learning in the shore crab (Carcinus maenas) is consistent with a key criterion for pain”, Journal of Experimental Biology, 216, pp. 353-358 [accessed on 25 December 2015].
Maldonado, H. & Miralto, A. (1982) “Effect of morphine and naloxone on a defensive response of the mantis shrimp (Squilla mantis), Journal of Comparative Physiology, 147, pp. 455-459.
Mather, J. A. (2001) “Animal suffering: An invertebrate perspective”, Journal of Applied Animal Welfare Science, 4, pp. 151-156.
Mather, J. A. (2008) “Cephalopod consciousness: Behavioral evidence”, Consciousness and Cognition, 17, pp. 37-48.
Mather, J. A. & Anderson, R. C. (2007) “Ethics and invertebrates: A cephalopod perspective”, Diseases of Aquatic Organisms, 75, pp. 119-129.
Tomasik, B. (2013) “Speculations on population dynamics of bug suffering”, Essays on Reducing Suffering [accessed on 18 March 2017].
Tomasik, B. (2015) “The importance of insect suffering”, Essays on Reducing Suffering [accessed on 23 March 2017].
Tomasik, B. (2016) “Brain sizes and cognitive abilities of micrometazoans”, Essays on Reducing Suffering [accessed on 18 December 2016].
Tye, M. (2017) Tense bees and shell-shocked crabs: Are animals conscious?, New York: Oxford University Press.
Volpato, G. L. (2009) “Challenges in assessing fish welfare”, ILAR Journal, 50, pp. 329-337 [accessed on 30 May 2013].
Walters, E. T. & Moroz, L. L. (2009) “Molluscan memory of injury: Evolutionary insights into chronic pain and neurological disorders”, Brain, Behavior and Evolution, 74, pp. 206-218 [accesed on 22 September 2013].
Wilson, C. D.; Arnott, G. & Elwood, R. W. (2012) “Freshwater pearl mussels show plasticity of responses to different predation risks but also show consistent individual differences in responsiveness”, Behavioural Processes, 89, pp. 299-303.
Zullo, L. & Hochner, B. (2011) “A new perspective on the organization of an invertebrate brain”, Communicative & Integrative Biology, 4, pp. 26-29.
1 Braithwaite, V. A. (2010) Do fish feel pain?, Oxford: Oxford University Press. Sherwin, O. M. (2001) “Can invertebrates suffer? Or how robust is argument-by-analogy?”, Animal Welfare, 10, pp. 103-108. Sneddon, L. U.; Braithwaite, V. A. & Gentle, M. J. (2003) “Do fishes have nociceptors? Evidence for the evolution of a vertebrate sensory system”, Proceedings of the Royal Society of London, Series B, 270, pp. 1115-1121. Elwood, R. W.; Barr, S. & Patterson, L. (2009) “Pain and stress in crustaceans?”, Applied Animal Behaviour Science, 118, pp. 128-136.
2 Crook, R. J. & Walters, E. T. (2011) “Nociceptive behavior and physiology of molluscs: Animal welfare implications”, ILAR Journal, 52, pp. 185-195 [accessed on 15 October 2013].
3 Wigglesworth, V. B. (1980) “Do insects feel pain?”, Antenna, 4, pp. 8-9. Allen-Hermanson, S. (2008) “Insects and the problem of simple minds: Are bees natural zombies?”, Journal of Philosophy, 105, pp. 389-415.
4 Balderrama, N.; Díaz, H.; Sequeda, A.; Núñez, A. & Maldonado H. (1987) “Behavioral and pharmacological analysis of the stinging response in africanized and italian bees”, in Menzel, Randolf & Mercer, Alison R. (eds.) Neurobiology and behavior of honeybees, Berlin: Springer, p. 127. Núñez, J.; Almeida, L.; Balderrama, N. & Giurfa, M. (1997) “Alarm pheromone induces stress analgesia via an opioid system in the honeybee”, Physiology & Behaviour, 63, p. 78.
5 This is a central question when it comes to how positive and negative experiences are spread in nature that is asked in a groundbreaking work in the examination of the suffering of animals in nature, Ng, Y.-K. (1995) “Towards welfare biology: Evolutionary economics of animal consciousness and suffering”, Biology and Philosophy, 10, pp. 255-285.
6 Bear in mind that others mollusks, such as cephalopods, have totally different nervous systems which are much more complex.
7 Crook, R. J. & Walters, E. T. (2011) “Nociceptive behavior and physiology of molluscs: Animal welfare implications”, op. cit.
8 Smith, J. A. (1991) “A question of pain in invertebrates”, ILAR Journal, 33, pp. 25-31 [accessed on 20 October 2013]. Sonetti, D.; Mola, L.; Casares, F.; Bianchi, E.; Guarna, M. & Stefano, G. B. (1999) “Endogenous morphine levels increase in molluscan neural and immune tissues after physical trauma”, Brain Research, 835, pp. 137-147. Cadet, P.; Zhu, W.; Mantione, K. J.; Baggerman, G. & Stefano, G. B. (2002) “Cold stress alters Mytilus edulis pedal ganglia expression of μ opiate receptor transcripts determined by real-time RT-PCR and morphine levels”, Molecular Brain Research, 99, pp. 26-33.
9 Morton, B. (2001) “The evolution of eyes in the Bivalvia”, in Gibson, R. N.; Barnes, M. & Atkinson, R. J. A. (eds.) Oceanography and marine Biology: An annual review, vol. 39, London: Taylor & Francis, pp. 165-205. Morton, B. (2008) “The evolution of eyes in the Bivalvia: New insights”, American Malacological Bulletin, 26, pp. 35-45. Aberhan, M.; Nürnberg, S. & Kiessling, W. (2012) “Vision and the diversification of Phanerozoic marine invertebrates”, Paleobiology, 38, pp. 187-204. Malkowsky, Y.; Götze, M.-C. (2014) “Impact of habitat and life trait on character evolution of pallial eyes in Pectinidae (Mollusca: bivalvia)”, Organisms Diversity & Evolution, 14, pp. 173-185. Morton, B. & Puljas, S. (2015) “The ectopic compound ommatidium-like pallial eyes of three species of Mediterranean (Adriatic Sea) Glycymeris (Bivalvia: Arcoida). Decreasing visual acuity with increasing depth?”, Acta Zoologica, 97, pp. 464-474.
10 Kamenos, N. A.; Calosi, P. & Moore, P. G. (2006) “Substratum-mediated heart rate responses of an invertebrate to predation threat”, Animal Behaviour, 71, pp. 809-813.
11 Charifi, M.; Sow, M.; Ciret, P.; Benomar, S. & Massabuau, J.-C. (2017) “The sense of hearing in the Pacific oyster, Magallana gigas”, PLOS ONE, 12 (10) [accessed on 24 January 2018].