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Another comment about uncertainty monitoring: Central place foragers tend to spend extra time memorizing the visual landmarks around their nest at times, there is a recent paper on ants describing how this correlates to uncertainty in some detail. As an insect moves further from its nest the accuracy of its knowledge of the nest decreases (errors accumulate in its path integration), and there is evidence that the magnitude accumulated error influences which search strategy an ant will use if it gets lost.
I also have a feeling that insects will start to ignore a sensorimotor cue that provides starts to provide unreliable information. For instance, airflow and visual motion are usually correlated to movement direction and used to control parameters like flight speed. If wind is artificially manipulated such that it is no longer correlated to visual motion or flight speed (it should be random, not negatively correlated), then I think the insect would stop using it as a cue to control flight speed. I can’t find a reference for this quickly, but I can look further if it’s of interest. I recall something similar also occurs in the case where two cues are initially paired with a reward during associative conditioning but only one turns out to be consistently rewarded (the distractor is called a confound) - after a while a bee can learn to ignore the confound and increase its accuracy. Again I don’t have a reference at hand for this but could look later.
Gavin, if you have the time, I’d love to see references for those abilities. We’ve got another round of invertebrate posts coming out in mid-to-late July, and most if not all of your examples can be used to bolster the case that arthropods in general and insects in particular deserve a close look from the effective animal advocacy movement. Thanks for your contributions!
Hi Jason, I look a bit more into the idea of uncertainty modelling in both sensorimotor and learning activities. I must admit I couldn’t find much on learning to ignore completely random cues (maybe I picked this idea up at a talk or discussion, not a paper, or I can’t get the right search terms for it), but I did come across a few extra studies associated with sensory processing, navigation and foraging that you might be interested in.
In terms of sensorimotor learning, the idea of humans doing probabilistic weighting of sensory cues according to their reliability was proposed of by Wolpert quite recently. There aren’t many insect studies looking at this directly, although some have considered this indirectly as part of the study design. Other useful search terms are probabilistic/dynamic/bayesian/optimal/multisensory reweighting/integration, reliability, uncertainty, and this all links closely to adaptive motor control.
A basic example is the idea of neural correlates that works to determine when several different sensory measurements agree—this was found a long time ago in locust neurons which fire most strongly when visual motion (as seen by both ocelli and compound eyes) and air flow indicate the same direction of motion. More recently moths have been found to increase their visual integration time at lower light levels, which would allow them to see more accurately at the expense of moving slower. Aphids have also been found to respond to indicators of predators approaching by temporally correlating multiple sensory cues in the case where individual cues may be unreliable. The above are probably examples of systems that innately deal robustly with uncertain information. However, I once did a study where I waxed or amputated honeybee antenna, in both cases removing their main sensory perception of airflow. The honeybees reacted to airflow differently in both cases, and I (very speculatively) suggested this could be because the honeybees that still had intact antenna believed the information from it, while amputated honeybees then tried to use alternate cues (air flow on body hairs and legs for instance). I don’t think my study is confirmation honeybees do sensory re-weighting (it wasn’t intended to be), but such proof may already exist in a similar context using cue conflicts or ablations.
In terms of navigation, I also found studies claiming optimal usage of navigation cues by ants and Drosophila.
There are also studies looking at foraging decisions made by bumblebees, which suggests that they prefer flowers that provide consistent nectar rewards, and they change their visitation rate to flowers depending on how likely the flower is to provide a reward.
Finally, there is a study suggesting that individual ants assess their uncertainty when deciding how to contribute to colony level decisions.
Hope these are helpful references with regards to uncertain insects!
Thanks for the references, Gavin! You truly are an inexhaustible resource. The paper on uncertainty monitoring in ants looks particularly impressive and relevant. I hope to give it a full read later this week. The ability of eusocial insects to incorporate diverse streams of information into an integrated decision-making framework is, to my mind, decent evidence that they are conscious.
(Also, I’m getting a session timed out error on the aphid link.)
No worries! Yes, eusocial insects certainly are quite amazing creatures. There are actually studies looking at facultatively social bee species (whereby females can nest individually or in hives with multiple reproductive females) that suggest sociality leads to increase in brain volume. Besides cognitive demands, sociality also appears to lead to other things like increased hygiene and immune function prevent disease spread in a colony.
Actually, it could be interesting to include naked mole-rats as a vertebrate comparison specific to social insects in this study. I’m not really familiar with their biology but they are generally considered eusocial , particularly that there is division of reproductive labour that creates queen and worker castes within colonies. Maybe impressive feats seen in social insects also appear in mole-rats more than you would expect compared to normal rats? In fact, there are also eusocial species shrimps from the Synalpheus genus which would probably display different traits to the other groups of crustaceans you’re looking at.
I also updated the Aphid link, it should work now, but the link is below if it doesn’t.
https://academic.oup.com/beheco/article/25/3/627/2900485
My comments are certainly biased towards bees because of my background. I hope there are relevant examples available for other invertebrates groups, although it may be that a lot of these concepts have mostly been tested in Drosophila or eusocial insects.
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.