Some snippets from some peer reviewed articles:
1. While similarity of nervous systems to that of
humans is an important criterion used in judging
whether or not an animal might feel pain, is it really
adequate? Like human engineering, biology provides
numerous cases where the same problem has
been solved in different ways (e.g. flight in bats,
birds and insects). Specifying design criteria does
not specify the solution. For that reason, some
thought is needed when dealing with animals that
are massively different from humans.
What, then, can be said about a special-purpose
damage-avoidance system in invertebrates? Insects
have complex nervous systems and the fruit fly, for
instance, can associate odours with electric shock
and avoid such odours on subsequent occasions
(Dudai et al. 1976). Insects have no fibre system
equivalent to the pain fibres of mammals. Moreover,
they will continue normal behaviour after
severe injury or removal of body parts (Eisemann
et al. 1984). A locust will continue to feed while
it is itself being eaten by a mantis. These and
other examples are taken as evidence that insects
are different from vertebrates in not sensing
pain (Wigglesworth 1980), but it is as well to
remember that mammalian herbivores may also
graze seemingly normally after severe injury.
Twenty years ago it would have
been hard to find a scientist wondering whether
or not insects felt pain. Nowadays, a number do
(e.g. Wigglesworth 1980; Eisemann et al. 1984;
Kavaliers 1988).
(
Patrick Bateson, Assessment of pain in animals, Animal Behaviour, Volume 42, Issue 5, November 1991, Pages 827-839, ISSN 0003-3472, 10.1016/S0003-3472(05)80127-7.(http://www.sciencedi...003347205801277)
2. Avoiding dangerous situations, such as noxious heat or harmful tissue damage, is certainly most fundamental behavior of all animals. One can hardly imagine that any animal could survive in nature without innate reflective mechanisms to avoid such situations or even to defend itself against them. Such avoidance reactions usually start with the activation of somatosensory neurons covering the body surface that are responsive to repellent cues, such as heat or strong mechanical stimulation. In vertebrates, processing of these peripheral signals in the brain can ultimately cause the sensation of pain [1]. The fruitfly Drosophila melanogaster has frequently been used as a valuable model organism to dissect the neuronal basis of a variety of sensory processes, for example olfaction, gustation or sound detection [2] and [3], which often are surprisingly similar to those of vertebrates. But do insects feel pain? Certainly not in the sense higher mammals do, but it is certainly possible that they have sensory systems for detecting stimuli that can cause injury or damage, and that the basic mechanisms are evolutionarily conserved from fruitflies to mammals.
Biologists use the term ‘nociception’ [4] and [5] for the sensory detection of potentially harmful stimuli, to differentiate it from the psychologically loaded term ‘pain’ [1]. Likewise, a defensive behavior elicited by such stimuli can be called ‘nocifensive’ behavior. Such a nocifensive behavior can be observed in such a simple organism as a
Drosophila larva. When fruitfly larvae are stimulated with a heated probe, such as the fine tip of a soldering iron, or are gently pinched, they perform a rolling behavior which is clearly different from their typical peristaltic locomotion behavior
[6]. To investigate the mechanisms causing such a nocifensive behavior one can ask whether there are dedicated nociceptive neurons in fruit flies, similar to nociceptive neurons of vertebrates.
In Drosophila, two morphological types of somatosensory neuron have been described. Type I neurons terminate in a ciliated dendrite and are located in sensory bristles that sometimes can form complex organs, such as the Johnston's organ, the fly's ear. Type I neurons are diverse with respect to the stimuli they respond to, sound or gravity for example. In contrast, type II neurons are simpler, as they lack sensory cilia and extend their naked dendrites along epidermal cells of the body surface. In this respect, these neurons, which are also called multidendritic neurons, are similar to the nociceptive neurons of vertebrates [5].
A function of Drosophila multidendritic neurons as true nociceptors has been proposed after the exciting finding that they express a channel protein of the TRP family that is homologous to a temperature-sensitive channel (TRPA1/ANKTM1) expressed in vertebrate nociceptive neurons [6]. This channel protein opens at temperatures above 38°C, ultimately activating the sensory neuron. Interestingly, deletion of this channel in the mutant
painless results in a defect in the nocifensive rolling behavior in
Drosophila larvae
[6], suggesting that multidendritic neurons might act as nociceptors. Other possible explanations for this remained, however; for example, it has been suggested that multidendritic neurons might be proprioreceptors, and if so, their impairment could simply prevent the fruitfly performing motor actions properly
[7]. A further complication is that multidendritic neurons are diverse: four different classes can be distinguished by their different dendritic arborisation patterns, perhaps indicating they have different functions.
As reported recently in Current Biology, Hwang et al.[8] have addressed this question in a study which demonstrates once again how favourable
Drosophila is for tackling such issues. Most importantly, transgenes can be expressed in
Drosophila easily in a variety of neuronal populations of interest, in this case within different subpopulations of multidendritic neurons. By expressing tetanus toxin, a poison that abolishes chemical synaptic transmission, the authors were able to block the output selectively from morphologically distinct classes of multidendritic neurons. They found that blocking of only one single type — class IV neurons — strongly affects the putative nociceptive response evoked by heat or mechanical stimulation. These data demonstrate that class IV neurons, but not the other classes, are required for the initiation or performance of the nocifensive behavior. In addition, the authors have used a more recently developed technology to artificially activate neurons by illumination. Such photo-activation of neurons can be achieved by expressing the light-sensitive cation channel ‘Channelrhodopsin-2’ (ChR2)
[9]. Illumination of the neurons expressing ChR2 causes their depolarization and thus activation.
The efficacy of this technology in Drosophila larvae has been demonstrated already in the context of olfactory learning and memory [10]. If multidendritic neurons are really nociceptive and responsible for triggering the nocifensive rolling behavior, light-induced activation of just these neurons should cause the illusion of a harmful stimulus and elicit the rolling response. Interestingly, only light-induced activation of class IV neurons induced a nocifensive rolling behavior, whereas activation of the other classes of neurons caused an accordion-like contraction of the larvae. This experiment is not only fascinating because of the modern genetic tricks that allowed Hwang
et al.[8] to turn light into a ‘harmful’ stimulus. The experiment clearly demonstrates that activation of class IV neurons is sufficient to cause the nocifensive rolling behavior, whereas the other classes of multidendritic neurons might serve different functions in the context of coordinated locomotion.
One observation was puzzling, however: the larvae rolled more often towards the side from which the noxious stimulus came rather than away from it. To understand this seeming paradox, Hwang et al.[8]considered how such a behavior might have evolved. In nature, a serious threat for insect larvae are parasitoids, insects whose larvae feed from the body of other insects
[11].
Drosophila melanogaster has such an enemy in the parasitoid wasp
Leptopilina boulardi, whose females penetrate the larvae with their ovipositor and lay their eggs inside the larval body. Hwang
et al.[8] showed that indeed
Drosophila larvae can defend themselves against such wasp attacks by performing their rolling behavior. In a kung-fu-like fashion, the larva wraps the wasp's sting around its body, flips the attacking wasp through the air and onto its back, which gives the larva time to escape. This fascinating new study by Hwang
et al.[8] vividly illustrates that animal behavior often can be understood only if the context of the animal's natural ecology is taken into account.
(André Fiala, Neuroethology: A Neuronal Self-Defense Mechanism in Fly Larvae, Current Biology, Volume 18, Issue 3, 12 February 2008, Pages R116-R117, ISSN 0960-9822, 10.1016/j.cub.2007.11.054. (http://www.sciencedi...960982207023408)
3. Invertebrates are an expansive and diverse group of animals that have had little attention regarding
anesthesia and analgesia. Economic use, environmental awareness, laboratory research, and increasing
demand for invertebrates as pets has lead to a greater desire for knowledge for these animals in the
veterinary medical community. With the increasing number of animal welfare regulations, various
scientific studies have improved the overall knowledge of invertebrate medicine, but much more research
is required to fully understand anesthesia techniques in the different species treated by veterinarians.
Analgesia is a controversial and often neglected topic with invertebrates because of the common belief
that invertebrates do not feel pain. Recently, the idea that invertebrates do not feel pain has been
challenged with the discovery of nociceptive pathways similar to those in vertebrates. This article presents
a general overview of anesthetics and analgesics used in selective invertebrate taxa
(
Gregory A. Lewbart, Conny Mosley, Clinical Anesthesia and Analgesia in Invertebrates, Journal of Exotic Pet Medicine, Volume 21, Issue 1, January 2012, Pages 59-70, ISSN 1557-5063, 10.1053/j.jepm.2011.11.007.(http://www.sciencedi...557506311002242)