Thursday, May 9, 2013

Intelligence and Pain in Invertebrates (Can Invertebrates Appreciate the Pain They Feel?)

The Bug Dude presents The Skeptical Entomologist 




INTELLIGENCE AND PAIN IN INVERTEBRATES

CAN INVERTEBRATES APPRECIATE THE PAIN THEY FEEL?


INTRODUCTION

   It's a popular topic among philosophical circles, and it is becoming more of an issues in science circles as well - is the experience of pain limited to humans, or can it be found in other organisms? In this case, "other organisms" means terrestrial invertebrates; dragonflies, butterflies, leeches, spiders, centipedes, worms, and the like. Unlike philosophy, however, science gives us the unique ability to not just ponder the question, but to solve it empirically. It may be a bit off topic, but it is worth stating before we start: neuroscience is slowly displacing psychology, and with each advancement in neuroscience we know more about the way the nervous system functions. As a side effect of such advancements, perhaps the most prominent point of strain is the concept of free will; this point isn't being dredged up to make a singular point - it has a significant role to play when we examine the responses to pain in organisms.
    However, to get back on track, it must be stated that it is undeniable that all organisms respond to stimuli. Whatever organisms you poke - be they bacteria, archaea, plants, fungi, or animals - will ultimately have some sort of response. However, the problem of pain becomes infinitely more complex when the subject is animals, as one could argue that only animals could even begin to process pain as we would (as we are animals ourselves). The reason this is infinitely more complex is, not only do we know that animals are capable (at least, some are in theory) of feeling pain, but we as humans have put a name to the pain. It may seem a rather odd distinction to make, and surely it has served our species well in distinguishing ourselves from our lowly origins.
    As an entomologist and arachnologist, the area of pain I am most interested in is whether or not terrestrial arthropods feel pain (I will also extend such academic curiosity to the mollusks with which we share the land). This is tied not only with the definition of pain (concrete or abstract), but also with intelligence (can an arthropod appreciate the pain it feels?). All organisms respond to stimuli, and it appears as though all animals, or at least a wide variety of them, feel what I will term "concrete pain" in this essay - that is to say, they react to pain from an objective, interactive world. Since we know that insects at least respond to pain (a caterpillar will, for example, react differently to being nudged with a finger as opposed to pricked with a needle), the question is how such experiences in the minute world relate to intelligence. It is most certainly not an easy question to answer, and I have no doubt I will ultimately fail in any attempt to do so with this essay. In science, we are seldom completely certain, but we can make intelligent, ethical, and rational judgments based on the evidence and information we have on hand.

THE MOST BASIC OF PAINS


   Before any sort of progress can be made, or, indeed, even before we can begin to answer the question of whether or not terrestrial invertebrates experience pain, we have to first define what we mean when we say "pain". The problem of pain may be infinitely more complex for us as organisms with complex nervous systems because pain is not as simple for us as it is for, say, a tarantula. As a starting point, we should define pain in its most basic form (as that is where we will start if we wish to assign more complex pain to invertebrates). Dictionary.com defines pain as "1. physical suffering or distress, cause by illness or injury; 2. a distressing sensation" [1]. The evolutionary purpose of pain is, therefore, to inform the animal when it is taking damage; an organism cannot survive and reproduce its genes if it is maimed or killed before it has the opportunity to do so. Pain provides the organism incentive to keep itself from harm. In its most basic level, it serves as a kind of physiological smoke alarm [2]. (You can do a simple experiment at home to demonstrate this. Take the tip of a sharpened knife, and push it into your flesh. Before you break the skin, you'll feel at least some small magnitude of pain; if you don't, you should probably consult an expert). In this most basic sense, it is quite apparent that invertebrates are subject to pain, or at least to an equivalent of it: If you nudge a beetle, say, it may move a little bit; however, if you take the same beetle and damage it, it is more than likely going to move away at a greater pace. It would seem logical to assert that there is a difference between pain and simply feeling something, as there is with "higher" animals such as Homo sapiens
     

INTELLIGENCE IN ARTHROPODS

    If we are going to look into the concept of pain in invertebrates, we are going to have to look at the level of brain development. Brains are amazing things - all that we see, touch, feel, hear, or will ever think is, essentially, confined to the brain. True, our senses are triggered by objective experiences (being poked with a stick, smelling a flower, bumping into a wall, etc.), but our own experience of them is confined within the brain. Brain development itself seems to correlate to higher intelligence and understanding to such a degree that when we think of super-intelligent extraterrestrials, they are almost always drawn with oversize, bulging brains.
    While the size of the brain is important, it's not necessarily the size of the brain alone. A sperm whale (Physeter macrocephalus) has a much larger brain than that of a human, yet it is the human, and not the whale, which have developed a global society which is now stretching its mechanical hand into space. What's important is brain-to-body ratio. That simply means that the brain size compared to the rest of the animal is what counts, and not brain size alone. Not only does brain size count, but brain function also depends on the neuronal activity in the brain [3].
A look at the brains off two of the arthropods we shall examine: a spider (top), and bee (bottom).

SPIDERS

   In terms of invertebrates, intelligence (or, at the very least, behaviors which seem intelligent) are readily available. Lets look at spiders - some of the most intelligent spiders in the world today are jumping spiders, who have brains so large they actually extend into the legs   of the animal (that's up to 80% of body size!).[4] The most widely appreciated of the jumping spiders belong to the genus Portia, which (as luck would have it) prey on other spiders. But do they use their brains to outwit other spiders?
A Portia spider makes an excellent introductory case study.  (http://www.flickr.com/photos/weett/4346160438/)

    
    The hunting behaviors of Portia spiders are an excellent place to start looking at intelligence levels of invertebrates (the most intelligent and emotional of all invertebrates, the Cephalopods, will not be taken into account, because, for this article, we are focused on terrestrial invertebrates). Portia spiders have often been referred to as "eight legged cats" because of their hunting behaviors. They not only hunt in the open, but have been known to build prey catching webs, and have been observed invading the webs of other spiders [4]. In laboratory settings, the spiders have even been observed solving problems that involve invading webs of spiders which are, as far as can be determined, foreign to them; in the Phillipines, Portia spiders have learned to hunt spitting spiders form behind (spitting spiders themselves hunt jumping spiders). While this appears to be instinctive, it often serves as a jumping-off point (pun intended) for trial-and-error attempts, form which the spider will learn. 

Japanese scientists have discovered that jumping spiders may use "image defocus" 
to judge distance between them and their prey items. 


    Intelligent behavior is exhibited by many different species of jumping spiders. Portia spiders are not alone in the spider world in their ability for visual processing on par with those of cats (well known for their keen eyesight). Some species have been observed in laboratories to recognize and react to images on computer screens; and, as remarkable ability as it is, it isn't even a new discovery - such intelligent behaviors were first observed in the 1980s. In one case, a Portia spider even attacked the screen when it recognized another spider on screen.[5] 
A Portia spider attacks a virtual prey species in a laboratory. 

    Jumping spiders are also known to react to humans in such a way as to make them unique from other spiders. When intimidated by a finger, the zebra jumping spider (Salticus scenicus) doesn't simply run or show it's fangs, as are stereotypical among spiders. It moves according to the movements of the finger, keeping an eye on it. Yes, it will ultimately try to escape if it feels threatened, but it's first reaction is not simply a fight or flight response. It seems to be curious about what it is its facing. Curiosity, as it turns out, is well documented in jumping spiders; they seem to be curious about any new thing which enters their immediate environment. [6]
    As with most solitary animals of their size, it is difficult to determine between intelligence and instinct. Surely one can say that a jumping spider can learn from trial-and-error, much the same way more "complex" animals tend to learn. On the other hand, approaching a spitting spider from behind seems to be hard-wired into the Portia spiders' brain, so one would be hard pressed to call that true "intelligence". Likewise, attacking the screen (an experiment in wolf spiders showed that they reacted simularly to simulated images) when they discern either a prey item or a rival may not be definitive sign of intelligence - it may not be intelligent at all. Once again, it may be nothing more than instinct (it is highly likely that instinct is the explanation). Jumping spiders make a strong case that arthropods may have some degree of intelligence, but they're a far cry from exploring space anytime soon.
Video of a jumping spider hunting a honeybee. 
The perfect (violent) transition to our next arthropod example.

HONEYBEES

    Honey bees are among the most important pollinators in the world. The most interesting facets of these insects is not just their famous way of communication (through the dance floor) but also their ability to learn and remember colors. However, most people don't know about bees for that reason - they are famous because of colony collapse disorder (CCD), which has been the the subject of documentaries (The Silence of the Bees, Where Have the Bees Gone?) and movies (The Happening makes a strong reference to CCD). While that is an interesting and major course of entomological resources (involving genetically modified organisms, pesticides, and a boatload of rage), what is relevant here is the behaviors exhibited by bees.
    The unique thing about bees is that the bees responsible for foraging for food seem to spend their entire times learning.[6] This certainly seems to indicate at least some level of intelligence (remember, intelligence in this case is strictly defined as the ability to learn). One could argue that the ability for honeybees to successfully learn over their lifetime is a handy trick delivered by the blind eye of evolution, and it seems as though that would play a large part in this scenario. On the other hand, it also seems blatantly absurd: the foraging bees are reproducing their own particular genes. To understand this phenomenon, we must first look at the social structure of honeybees.
    Honeybee society revolves around the queen bee, the largest member of the hive which produces the progeny which ultimately constitute the entirety of a hive, and can live for three to five years. Queen bees are develop with the death of an existing queen, in which case a larvae (the vast majority of bees are female) are fed royal jelly.[7] (An interesting side note - in many hives which are affected with CCD, the queen bees are often still present) Once in her life, the queen will mate with a drone bee, the only male bees present in the entirety of the hive. Drones are stouter bees which are larger than the workers but smaller than the queen, with larger eyes and are unable to sting (bee stingers are modified ovipositers, which are understandably absent in male bees); it is worth noting that drone bees are haploid. Finally, the vast majority of the hive are sterile, female daughters of the queen, who perform food gathering and defense roles in the hive.
    Having reviewed the social structure of the honeybee, we can now begin to determine whether the remarkable abilities they exhibit are intelligent or if they are merely instinctual. If they are intelligent, we have to determine if such behaviors are taught or arrived at from trial and error.
    First, lets look at the ability of bees to learn over the course of their life. Evidence suggests that this may be intelligence; memory formed from a single experience can last for up to three days, and when it is enforced three or more times, it lasts for the bee's lifetime.[8] It's quite an amazing thing to think about; that an animal so many of us overlook, that so many of us may even despise, is capable of learning and remembering experiences. To further illustrate this point: in one experiment, bees were exposed to a variety of colors, one of which was attached to a sugary reward. After one trial, the bees were kept in a cage for several days. When the experiment was finally repeated a few days later, over 50% of the bees began foraging with the correct color (the one that had been associated with the sugary treat).[9] 
     Is the communication through dance intelligence? If we assert that bees are intelligent, as experiments suggest they are, are the complex physical communications intelligent as well? Two hypotheses for how bees communicate the location of productive foraging areas are as follows:

1: Honeybees "explain" the location of productive foraging areas through body movements
2: Bees use chemical signals to "explain" the location of productive foraging areas.

    By far the more accepted of the two ideas is the dance-communication proposal, which, in the interest of brevity, will be explained in the video below:

A video explaining an interesting experiment with dance-communication of honeybees. 
Karl von Frisch eventually won a Nobel Prize in Physiology or Medicine for his efforts.

    But is the dancing communication of honeybees instinctive or learned? As it turns out (most non-entomologist or insect enthusiasts would not be aware of this), the dance communication (also known as the waggle dance, or Tanzsprache, literally "dance-talk", as defined by von Frisch) is rather controversial. It is still debated whether or not it counts as actual language, which would play a major role in whether or not it is intelligence or instinct. Tanzsprache seems to signify something (the location of productive foraging areas) through a signifier (the dance itself), it lacks any form of symbols, syntax, or grammar.[10] As it seems to be only the basic possible form of communication, however amazing for arthropods, one can't help but feel that it is deeply instinctual. This is only confirmed even more when it is taken into consideration that all bee species have a Tanzsprache, although, much like human language, there seem to be different variances depending on species (in humans, the differences are due to geographical location). It is interesting to note that there is variability among the successfullness of the Tanzsprache - some honeybees can successfully find the foraging area after watching less than ten repetitions of the dance, while others seem to be unable to find it after watching over fifty.[10] 
Bees can memorise [sic] at least six locaitons, and three paths leading to each. They can remember at least four good choices and four bad choices. These memories are not just visual; they can also remember them by smell, and the overall memory is often a mixture of smell, location and colour."[13]
    Honeybees are also an intriguing species because they seem to display, at least on some level, emotions. When exposed to negative circumstances, the bees displayed negative cognitive biases:

"Shaken bees also have lower levels of hemolyph dopamine, octopamine, and serotonin. In demonstrating state-dependent modulation of categorization in bees, and thereby a cognitive component of emotion, we show that the bees' response to a negatively valenced  [sic] event has more in common with that of vertebrates than previously thought."[15] 

DRAGONFLIES

    I previously mentioned dragonflies in my last Skeptical Entomologist post, in which I looked at homosexuality among several species. However, what is even more amazing (and, for us human "elitists" so sure of our superiority over insects, intimidating) is the recent discovery that dragonflies have the ability to engage in what is termed "selective attention". That means that, if the dragonfly is following a group of midges, for example, the predator will have the ability to focus on one individual within the swirling mass. This raises the interesting question, because the ability to focus on one thing in particular in a chaotic environment is definitely something we have long thought only occurred in animals with higher cephalization and brain-to-body ratios.
    A discovery of incredible importance in cognitive research and with applications in robotics, this arguably major announcement was made in 2012.[11] And yes, while it was slightly overshadowed by the discovery of the Higgs boson (which was a long and time consuming process, so perhaps the discovery of the Higgs deserved the spotlight), that fact does not negate its importance. The truly unique aspect of this discovery was not just that the behavior was observed in insects, but that the process by which it was possible was also discovered: the neural activity. In fact, it was the first neural activity which allowed for selective attention in any invertebrate.[11] This ability allows the dragonfly to hunt small insects - such as mosquitoes and midges - with a 97% success rate.
Dr. Steven Wiederman, one of the scientists who
 discovered the ability of selective attention in dragonflies. 
(Photo credit: David O'Carroll, University of Adelaide) 

    The dragonfly case is interesting because it further supports the assertion that the behaviors, personality, and ultimate overall character of an animal is dependent on brain function. While there may be naysayers like Deepak Chopra who will ultimately scream in the face of anybody that will listen ("The brain and the mind exist separately" is one of Chopra's most famous, and unsupported, claims), that does not detract from the essence of such a discovery. Given current knowledge about neural pathways and brain function, it is very reasonable to assume that the dragonfly's success rate at hunting would plummet horribly if this particular neural pathway was interrupted.


   DO ARTHROPODS EXHIBIT COGNITIVE ABILITIES?

    As with intelligence, the answer largely depends on how the word "cognitive" is defined. According to dictionary.com, the first definition of cognition is "1. or or pertaining to the act or process of konwing, perceiving, remembering, etc.; of or relating to cognition; 2. of or pertaining to the mental processes of perception, memory, judgement, and reasoning, as contrasted with emotional and volitional processes."[12] As the initial definition is very much in alignment with the definition of intelligence, we will use the second definition for the purpose of this essay. Can insects and spiders reason? Can they make judgments? In my opinion, the answer is decidedly yes. Spiders, such as the jumping spiders, assess situations before they attack their prey, looking for the best possible angle which maximizes jumping potential and minimizes potential injuries. If insects and spiders are so different from us, yet display at least some cognitive functions, what does that mean for their experience of pain? 
    Do insects think? If they do, it is probably not in a manner to which human would consider thinking. Higher brains provide the ability to think abstract thoughts. An insect or spider may be said to "think" if we wish to define thinking as only the processes of the brain. They certainly seem to be able to assess situations and act accordingly.
    


DEFINING THE HYPOTHESES 

    Now that we have reached the main topic of discourse for this essay - Do insects appreciate (understand) the pain they feel? - it is time to take a short break and decide on our pair of hypotheses. Take a break, get up and walk around, have a shot of Dry Fly or Buffalo Trace, and come back when you're ready.

    Ready? Good. We can now define the hypotheses we wish to test:
Null hypothesis: arthropods do not understand/appreciate the pain they experience.
Alternative hypothesis: arthropods do understand/appreciate the pain they experience.

     Now that that is settled, we may continue our discussion. First, we should start off with what we as humans experience as emotional or abstract pain, and why our illusion of free will may be clouding our judgement on the issue. 


HOW "FREE WILL" CLOUDS THE DEBATE

    If we ever wish to determine the answer to the question of arthropods understanding their pain, we must first understand what our pain is and how we, as humans, understand it. Having a much more complex social structure which allows for the spreading of memes, a higher degree of cognitive capabilities, and a far more developed sense of individual identity, our own appreciation of our pain is bound to be vastly different than that of any arthropod. Or so you would think. What if I were to propose to you that our experience of pain is, evolutionary  almost exactly the same? What if all of our cultural trappings, religious traditions, and ever-present memes were, at their very core, completely wrong? What if we're not different from the animal kingdom?
    I propose that the problem of human pain is one of the main ways our species has separated itself from the rest of the organic world - in our drive to single ourselves out as divinely special, we attributed pain only to ourselves. When it became overwhelmingly apparent that other animals feel physical pain as well, our species desperately clung to the meme that our species could experience emotional and abstract pain. As is usual when you learn more about the natural world, that idea was also smashed. Other animals, such as chimpanzees and elephants, experience emotional pain; this can be seen best when a member of their close family dies. Eventually, we came to understand it, thanks to the enthusiasm of amateur and professional biologists alike, keeping the intellectual curiosity alive.
    It seems as though the idea of insects understanding pain strikes many people even deeper. "At least," they seem to think to themselves, "at least we are different from insects! How far removed are we from the ant, and that has made us so superior!" Except, of course, that the ant has been evolving over the eons a well. As humans, we remain only partly rational, and very reluctant to accept new ideas (in fact, one of the basic tenants of scientific literacy is the replacement of private prejudice with accepting ideas based on publicly verifiable evidence) when they pop up. Our species even suffers from cognitive dissonance, in which two contradictory beliefs are held by the same individual (that is, of course, a highly simplified definition, which I will tackle in a later post).


HOW "FREE WILL" HAS FOOLED US ALL

    Before we begin, I should preface this by saying that, while free will is an illusion (or, as neuroscientist Sam Harris points out, an illusion of an illusion), we all are autonomous; that is, just because we don't have the free will we've been tricked into believing we have does not make us puppets. It simply makes us mistaken. If you wish to know more, I recommend reading Free Will by Sam Harris. 
    The concept of free will has been one of the defining characteristics of mankind; it shapes our justice system, our religions, are thoughts on personal responsibility. Essentially, this idea that we are each the conscious authors of our own actions is perhaps man's greatest intellectual lie. As I have mentioned in the beginning of this essay, psychology has been replaced with neuroscience. We can now look into the brain and understand what is going on, understand how it works, and map it. And, once again, a comfortable myth has been shattered: neuroscience is proving us that each of our individual actions are made subconsciously seven seconds before we become aware of any intent to perform said action. The famous example is the subject in a laboratory, who is asked by scientists to raise one of his hands at random; the observing scientists can not only tell when he is going to raise his hand, they can predetermine what hand he will raise. 
    This begs the question - if we are not the original authors of our thoughts, what are we? The mind - the part of the brain that we call ourselves, the conscious part - may be nothing more than a byproduct of an evolving brain; or, perhaps, it is the next step and consciousness is the direct product of a more complex brain. Whatever the initial cause, the fact remains the same - we no more consciously choose our own actions than a cricket may choose it's actions. 
    What does this mean? This means, of course, that we are essentially animals of instinct. We are doing what our evolution has allowed us to do. And while it may be much more complex than the smartest Portia spider (after all, Portia spiders don't build drive Porches), the argument can be made that it is essentially the same. After all, if we are simply observers of our own lives, watching as it plays out, than we are no more in control than the spider. 
    But, I stress again, we have deceived ourselves. We think that we are the conscious authors of our own lives, and we see arthropods as mindless, busy creatures who do nothing more than play their part. In truth, that is all humans do, although humans have the happy little ability of changing their behaviors (thanks to our highly evolved brain).
I'll just let Sam Harris himself explain the illusion of free will.

CONCLUSION 

    The answer seems to be a resounding yes. If we understand the fact that brain functions lead to behaviors, actions, emotions, and, ultimately, consciousness, then we have no other choice but to acknowledge that other animals capable of expressing anything similar to those have the ability to understand their pain. Many may find such an idea unbearably uncomfortable, but, as skeptics, it is something we have to consider. We cannot be closed minded to evidence, even if it suggests a reality contrary to what we wish reality was. Allow me to give an example: when I was a child, I used to butcher bees with one of my first childhood friends. The very fact that there is evidence which indicates they are capable of even the most rudimentary emotions leaves me utterly horrified by my past actions, leaving me feeling like quite a monster. I'm certain I am not the only one who has done such things (if it's any consolation, at the time we didn't know any better; we were ignorant of reality). But the past is the past, and we must reconcile ourselves with reality, even if it means looking back at what was (to us) a fun summer afternoon, even if it takes on a hue of brutal mania.
    To quote a study which appeared in the journal Animal Welfare,

"By closely examining the responses of invertebrates, it can be seen that they often behave in a strikingly analogous manner to vertebrates.... invertebrates such as cockroaches, flies and slugs have short- and long-term memory; have age effects on memory; have complex spatial, associative and social learning; perform appropriately in preference tests and consumer demand studies; exhibit behavioural and physiological responses indicative of pain; and, apparently, experience learned helplessness. The similarity of these responses to those of vertebrates may indicate a level of consciousness or suffering that is not normally attributed to invertebrates. This indicates that we should either be more cautious when using argument-by-analogy, or remain open-minded to the possibility that invertebrates are capable of suffering in a similar way to vertebrates."[14]
    Such a quote makes one stop to wonder. Perhaps we feel pain the exact same way an insect or spider does, but maybe the difference is that our species has the ability to ponder the pain and, ultimately, stop it. Does this knowledge mean that we can't kill insects which may be dangerous to ourselves or our families? Of course it doesn't; however, it does strongly suggest that we find a more reasonable solution than simply smashing the life out of a spider. Not only is it a crucial member of the local food web, if you squash it but fail to kill it, it may suffer.
    As animals as complex as ourselves, we need to act in an intelligent manner. This is hard, as our species is naturally a violent, xenophobic species that usually displays high levels of megalomania, typically seeing anything less than it as inferior. Even though the answer to the arthropod/pain question is important and certainly interesting, our behaviors toward them should remain the same in either case. As intelligent mammals capable of empathy, it is best to have the mental approach of utilitarianism - do the least amount of harm as possible, and do the most good as possible.
   

SUGGESTED READING

(1) http://dictionary.reference.com/browse/pain
(2) http://www.socrethics.com/Folder2/Biology.htm#C3
(3) http://curiosity.discovery.com/question/is-bigger-brain-better
(4) http://www.informatics.sussex.ac.uk/research/groups/ccnr/Papers/Downloads/Harland_Cimb2000.pdf
(5) http://www.cabinetmagazine.org/issues/25/wertheim.php
(6) http://psych.mcmaster.ca/dukas/Dukas%20%26%20Visscher%2094.pdf
(7) http://www.backyardbeekeepers.com/facts.html
(8) http://www.jneurosci.org/content/15/3/1617.full.pdf
(9) http://en.wikipedia.org/wiki/Bee_learning_and_communication#cite_note-Menzel-4
(10) http://en.wikipedia.org/wiki/Waggle_dance#cite_note-GF2009-10
(11) http://www.sciencedaily.com/releases/2012/12/121220143224.htm
(12) http://dictionary.reference.com/browse/cognitive
(13) http://bioteaching.wordpress.com/2010/05/03/insect-brains-and-animal-intelligence/
(14) http://www.scopus.com/record/display.url?eid=2-s2.0-0001460737&origin=inward&txGid=Fefoa6RxV0qF3gMVRibiJw3%3A2
(15) http://www.cell.com/current-biology/abstract/S0960-9822%2811%2900544-6

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