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.
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.
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.