A few thoughts about the categories in this article:
-Deception: There are some species of cuckoo bees that will sneak into the hive of another (in this case a solitary) bee, eat the owners eggs and then lay there own. As is the case with cuckoo birds, the owner then happily raises them as her own.
More extreme cases of nest parasitism occur in bumblebees when a cuckoo bumblebee invades a newly established hive of a true bumblebee, kills its queen, and then uses the original queen’s workers to raise her own offspring (the cuckoo bumblebee can only lay fertilised eggs, not workers). The later is more complex than a passive act of deception, although it’s also not clear to what extent the original workers are completely deceived or just being dominated by the invader.
-Self-control: I’m not sure that comparing self-control between feeding and reproductive contexts is really appropriate. Maybe a better choice would be fungus gardening or aphid herding by ants: In the former case the ants don’t eat the leaves they collect in order to grow fungus on them (although I am not sure the ants could actually digest the raw leaves), in the later case they don’t eat the aphids so they can milk them (this needs a video).
The self-control of bees is kind of imposed by most workers being sterile and the queen dominating them. This is also not universally true, and the weird relatedness between bee colony members and the occasional presence of workers with developed ovaries mean that it is advantageous for workers to lay male eggs if they have the opportunity (unfertilized honeybee eggs produce male clones of their mother; so a bee is most related to her sons, potentially more related to her sisters ((if they have the same father)) and their sons than her mother, and least related to her brothers—I’m not sure this is true for all social bee species). In bumblebees this can result in a worker revolts where the workers in an established colony kill their queen and all start laying male eggs.
-Paying a cost to receive a reward: Aphid herding ants defend their aphids from predators/competitors and it seems that they make a cost-benefit type decision about if they will defend them.
-Tool use: I think that prolonged nest construction kind of fits in here. External resources need to be collected over time (different bees use combinations of mud, resin, cotton, flower petals, small rocks, and other items to build their nests) in specific sequences, the cost is lost time foraging for food, and the benefit of the nest might not be realized until it is finished (or gained progressively during construction, it’s not useful straight away like the hermit-crab’s shell).
Thanks for the examples; keep them coming! Whether or not they possess the capacity for valenced experience, eusocial insects truly are remarkable creatures. Do you have an easy reference for the cuckoo bumblebee behavior? I’ve got a running list of amazing things different invertebrates do, and I’d love to add it to the list.
(On the subject of videos, check out the video I’ve linked in footnote 53. It always brings a smile to my face.)
To extend the tool use point a bit, I recall that primates have been found to have extra neurons in sensorimotor brain regions that are most active when the animal is using a tool, and essentially provide extra capacity for the brain to extend sensory and motor mappings/homunculus to include external artifacts (apparently also quite useful when learning to control of things with BCI). I’m not sure if this type of latent neural capacity has been found in rodents and strongly suspect it wouldn’t be present in insects (they tend to be quite frugal with their neurons!), although tool using birds like crows may have been studied as a comparison. Having neural circrity for tool use should be a sufficient (but perhaps not necessary) criteria for flexible tool use and its quite an objective (if difficult) test.
I read this in Beyond Boundaries by Miguel Nicolelis (good book although a bit long winded and fanciful) which should have some academic references.
Actually, Nicolelis’s BCI work also has some relevance to self-recognition. You can put electrodes into a monkey’s motor cortex, measure the neural activation associated with, say, arm movement and then decode those signals to control the motion of a robot arm (that the monkey is is not aware of) pretty well. However, if you show the monkey the arm and it is rewarded for moving the robotic arm, it often stops moving its own arm while continuing to use the disembodied arm (with pretty much the same motor cortex activity). I’d never thought of this in the context of awareness before, but suggests it is somewhat analogous to a mirror test and overcomes some of the limitations you mentioned. A fair bit of a work has been done around insect neural interfaces (probably more invasive and extreme than anything an ethics board would let you do to a mammal to be honest) and you might find that similar tests have been performed but not labeled as a self-recognition tests.
Thanks Gavin! I’ve added Beyond Boundaries to my reading list.
The potential connection between BCI and self-recognition is fascinating. Offhand, do you know any references for insect neural interface studies that might be comparable to the monkey example you describe?
The example that first springs to mind is the work of Kanzaki’s group who study odour plume tracking in silk moths. They have made robots controlled by both a moths walking action (also a movie) and also by its measured neural activity. However, when doing electrophysiology on insects it is common to completely wax their body in place and amputate their legs/wings to minimize electrical noise caused by muscle movement (which they did in the moth case). I’d forgotten this, and it does make it a bit harder for insects to demonstrate self-awareness in a similar way to the monkeys. Still, it’s recently become more common to make recordings from actively behaving insects, as active behaviour has been found to modulate many neural responses (such as optic lobe processing of visual motion), so some more relevant examples might have been published recently.
A few thoughts about the categories in this article:
-Deception: There are some species of cuckoo bees that will sneak into the hive of another (in this case a solitary) bee, eat the owners eggs and then lay there own. As is the case with cuckoo birds, the owner then happily raises them as her own.
More extreme cases of nest parasitism occur in bumblebees when a cuckoo bumblebee invades a newly established hive of a true bumblebee, kills its queen, and then uses the original queen’s workers to raise her own offspring (the cuckoo bumblebee can only lay fertilised eggs, not workers). The later is more complex than a passive act of deception, although it’s also not clear to what extent the original workers are completely deceived or just being dominated by the invader.
-Self-control: I’m not sure that comparing self-control between feeding and reproductive contexts is really appropriate. Maybe a better choice would be fungus gardening or aphid herding by ants: In the former case the ants don’t eat the leaves they collect in order to grow fungus on them (although I am not sure the ants could actually digest the raw leaves), in the later case they don’t eat the aphids so they can milk them (this needs a video).
The self-control of bees is kind of imposed by most workers being sterile and the queen dominating them. This is also not universally true, and the weird relatedness between bee colony members and the occasional presence of workers with developed ovaries mean that it is advantageous for workers to lay male eggs if they have the opportunity (unfertilized honeybee eggs produce male clones of their mother; so a bee is most related to her sons, potentially more related to her sisters ((if they have the same father)) and their sons than her mother, and least related to her brothers—I’m not sure this is true for all social bee species). In bumblebees this can result in a worker revolts where the workers in an established colony kill their queen and all start laying male eggs.
-Paying a cost to receive a reward: Aphid herding ants defend their aphids from predators/competitors and it seems that they make a cost-benefit type decision about if they will defend them.
-Tool use: I think that prolonged nest construction kind of fits in here. External resources need to be collected over time (different bees use combinations of mud, resin, cotton, flower petals, small rocks, and other items to build their nests) in specific sequences, the cost is lost time foraging for food, and the benefit of the nest might not be realized until it is finished (or gained progressively during construction, it’s not useful straight away like the hermit-crab’s shell).
Hi Gavin,
Thanks for the examples; keep them coming! Whether or not they possess the capacity for valenced experience, eusocial insects truly are remarkable creatures. Do you have an easy reference for the cuckoo bumblebee behavior? I’ve got a running list of amazing things different invertebrates do, and I’d love to add it to the list.
(On the subject of videos, check out the video I’ve linked in footnote 53. It always brings a smile to my face.)
No worries Jason, happy to keep posting the examples that come to mind (finally my knowledge of obscure insect behaviours is useful in EA!). This is a recent review of bumblebee cuckoos that could be useful. I also found another study indicating bumblebee cuckoos actively change their odor profiles to maintain control over the hives workers.
I agree, bumblebees look amazingly cute when rolling balls around! The string pulling experiment done by the same lab also has a nice video.
To extend the tool use point a bit, I recall that primates have been found to have extra neurons in sensorimotor brain regions that are most active when the animal is using a tool, and essentially provide extra capacity for the brain to extend sensory and motor mappings/homunculus to include external artifacts (apparently also quite useful when learning to control of things with BCI). I’m not sure if this type of latent neural capacity has been found in rodents and strongly suspect it wouldn’t be present in insects (they tend to be quite frugal with their neurons!), although tool using birds like crows may have been studied as a comparison. Having neural circrity for tool use should be a sufficient (but perhaps not necessary) criteria for flexible tool use and its quite an objective (if difficult) test.
I read this in Beyond Boundaries by Miguel Nicolelis (good book although a bit long winded and fanciful) which should have some academic references.
Actually, Nicolelis’s BCI work also has some relevance to self-recognition. You can put electrodes into a monkey’s motor cortex, measure the neural activation associated with, say, arm movement and then decode those signals to control the motion of a robot arm (that the monkey is is not aware of) pretty well. However, if you show the monkey the arm and it is rewarded for moving the robotic arm, it often stops moving its own arm while continuing to use the disembodied arm (with pretty much the same motor cortex activity). I’d never thought of this in the context of awareness before, but suggests it is somewhat analogous to a mirror test and overcomes some of the limitations you mentioned. A fair bit of a work has been done around insect neural interfaces (probably more invasive and extreme than anything an ethics board would let you do to a mammal to be honest) and you might find that similar tests have been performed but not labeled as a self-recognition tests.
Thanks Gavin! I’ve added Beyond Boundaries to my reading list.
The potential connection between BCI and self-recognition is fascinating. Offhand, do you know any references for insect neural interface studies that might be comparable to the monkey example you describe?
The example that first springs to mind is the work of Kanzaki’s group who study odour plume tracking in silk moths. They have made robots controlled by both a moths walking action (also a movie) and also by its measured neural activity. However, when doing electrophysiology on insects it is common to completely wax their body in place and amputate their legs/wings to minimize electrical noise caused by muscle movement (which they did in the moth case). I’d forgotten this, and it does make it a bit harder for insects to demonstrate self-awareness in a similar way to the monkeys. Still, it’s recently become more common to make recordings from actively behaving insects, as active behaviour has been found to modulate many neural responses (such as optic lobe processing of visual motion), so some more relevant examples might have been published recently.