The Conscious Nematode: Exploring Hallmarks of Minimal Phenomenal Consciousness in Caenorhabditis Elegans

By Vasco GrilošŸ”ø @ 2025-12-05T17:35 (+40)

This is a linkpost to https://pmc.ncbi.nlm.nih.gov/articles/PMC10723751/

This is a linkpost for The Conscious Nematode: Exploring Hallmarks of Minimal Phenomenal Consciousness in Caenorhabditis Elegans by Diego Becerra, Andrea Calixto, and Patricio Orio, which was originally published in the International Journal of Psychological Research in October 2023. Below are the abstract, summary of the evidence for consciousness in Caenorhabditis (C.) elegans, and my quick thoughts.

Abstract

While subcellular components of cognition and affectivity that involve the interaction between experience, environment, and physiology -such as learning, trauma, or emotion- are being identified, the physical mechanisms of phenomenal consciousness remain more elusive. We are interested in exploring whether ancient, simpler organisms such as nematodes have minimal consciousness. Is there something that feels like to be a worm? Or are worms blind machines? ā€˜Simpler’ models allow us to simultaneously extract data from multiple levels such as slow and fast neural dynamics, structural connectivity, molecular dynamics, behavior, decision making, etc., and thus, to test predictions of the current frameworks in dispute. In the present critical review, we summarize the current models of consciousness in order to reassess in light of the new evidence whether Caenorhabditis elegans, a nematode with a nervous system composed of 302 neurons, has minimal consciousness. We also suggest empirical paths to further advance consciousness research using C. elegans.

Summary of the evidence for consciousness in Caenorhabditis elegans

Table 1. Summary of the Current Evidence of the Criteria Reviewed in the Literature for Experiencing (Phenomenal Consciousness) in C. Elegans.

CriteriaFound in C. elegansReferences
Differentiated states and global activityYes. C. elegans forward and backward gaits are generated by either a reflex chain, or one or more CPG’s [central pattern generators]. Also, pirouettes and other motor commands require reverberating collective nested neural dynamics. Finally, it is argued that C. elegans have true sleep states, and its behavior changes globally under some anesthetics.Wen et al., 2012Olivares et al., 2018Kato et al., 2015Kaplan et al., 2020Trojanowski & Raizen, 2016Nichols et al., 2017Nambyiah & Brown, 2021
Binding and integrationYes. There is evidence for multisensory (chemical, mechanical and thermal) integration. Also, several sensory neurons in C. elegans are polymodal. Finally, a computational model of C. elegans’ neural activity show positive values of Phi (Ļ•) [see integrated information theory]Ghosh et al., 2017Metaxakis et al., 2018Goodman & Sengupta, 2019Antonopoulos et al., 2016
Flexible evaluative systemPerhaps. C. elegans is capable of non-associative and associative learning, and of integrating cues with opposing value in order to make a decision. There is some evidence of experience-dependent preferences, gustatory plasticity, and second order learning.Ardiel & Rankin, 2010Ghosh et al., 2017Guillermin et al., 2017Dekkers et al., 2021Merritt et al., 2019
Embodiment and selfPerhaps. There is an efferent copy of motor commands. Communication with bacteria promotes intergenerational memories depending on the bacterium’s virulence. Induction of diapause is pheromone-dependent and thus, C. elegans can recognize its community.Ji et al., 2021Palominos et al., 2017Gabaldon & Calixto, 2019
IntentionalityNot known. Mapping of worldly objects or C. elegans’ own body in neuronal networks seems limited, and goaloriented behavior doesn’t seem to be informed by internal states decoupled from the immediate environment. 
Selective attentionNot known. 

My quick thoughts

I think further research on the sentience of nematodes would be useful. This is one of the ā€œFour Investigation Prioritiesā€ mentioned in section 13.4 of chapter 13 of the book The Edge of Sentience by Jonathan Birch. However, I believe investigating the (expected hedonistic) welfare per animal-year of nematodes and other soil animals conditional on sentience is a much higher priority. I am much more uncertain about it than the probability of sentience. I would not be surprised if I changed my view that one should optimise for increasing the welfare of soil animals and maybe microorganisms. Meanwhile, I continue to recommend research informing how to increase the welfare of soil animals.


huw @ 2025-12-06T08:44 (+6)

Small drive-by question for you: In your opinion, if C. Elegans is conscious and has some moral significance, and suppose we could hypothetically train artificial neural networks to simulate a C. Elegans, would the resulting simulation have moral significance?

If so, what other consequences flow from this—do image recognition networks running on my phone have moral significance? Do LLMs? Are we already torturing billions of digital minds?

If not, what special sauce does C. Elegans have that an artificial neural network does not? (If you’re not sure, where do you think it might lie?)

(Asking out of genuine curiosity—haven’t had a lot of time to interface with this stuff)

Vasco GrilošŸ”ø @ 2025-12-06T10:25 (+6)

Thanks for the questions, Huw!

In your opinion, if C. Elegans is conscious and has some moral significance, and suppose we could hypothetically train artificial neural networks to simulate a C. Elegans, would the resulting simulation have moral significance?

I would say the moral significance, which for me is the expected hedonistic welfare per unit time, of the simulation would tend to that of the C. Elegans as more components of this were accurately simulated. I do not think perfectly simulating the behaviour is enough for the moral significance of the simulation to match that of the C. Elegans. I believe simulating some of the underlying mechanisms that produced the behaviour may also be relevant, as Anil Seth discussed on The 80,000 Hours Podcast.

Consciousness does not necessarily imply valenced (positive or negative) subjective experiences (sentience), which is what I care about (I strongly endorse hedonism). C. Elegans being conscious with 100 % probability would update me towards them having a greater probability of being sentient, but not that much. I am mostly uncertain about their expected hedonistic welfare per unit time conditional on sentience, not about their probability of sentience. I would say everything, including a Planck volume in deep space vacuum, could have a probability of sentience of more than, for example, 1 % if it is operationalised in a very inclusive way. However, more inclusive operationalisations of sentience will lead to a smaller expected hedonistic welfare per unit time conditional on sentience. So I would like discussions of moral significance to focus on the expected hedonistic welfare per unit time instead of just the probability of sentience, or just the expected hedonistic welfare per unit time conditional on sentience.

If so, what other consequences flow from this—do image recognition networks running on my phone have moral significance? Do LLMs? Are we already torturing billions of digital minds?

I think increasing the welfare of soil animals will remain much more cost-effective than increasing digital welfare. Assuming digital welfare per FLOP is equal to the welfare per FLOP of a fully healthy human, I calculate the price-performance of digital system has to surpass 2.23*10^27 FLOP/$ for increasing digital welfare to be more cost-effective than increasing the welfare of soil animals, which corresponds to doubling more than 29.0 times starting from the highest one on 9 November 2023. One would need 60.9 years for this to happen for Epoch AI’s doubling time of the FP32 price-performance of machine learning (ML) hardware from 2006 to 2023 of 2.1 years.

Elijah Whipple @ 2025-12-17T17:34 (+3)

Hey! Are you aware of Elizabeth Irvine's work? She has a paper where she talks about how C. Elegans is plausibly not conscious, but meets a lot of the current criteria people use for consciousness. She uses this as a way to critique these criteria. 
I tend to agree with her and think that the criteria people use for consciousness are pretty made up. I think consciousness research could be moved forward with methods like the ones from this paper. (I think Birch wrote something that exposed me to this paper.)

Vasco GrilošŸ”ø @ 2025-12-17T19:24 (+4)

Thanks for comment, Elijah!

I did not find a free version of Elizabeth's article Developing Valid Behavioral Indicators of Animal Pain, but I have now read this summary from Gemini, and this related article by Elizabeth. I agree with her that nociception does not necessarily imply sentience, but I do not rule this out. Here are some paragraphs I like from the article ā€œAll animals are consciousā€: Shifting the null hypothesis in consciousness science by Kristin Andrews which relate to the article from Elizabeth you shared.

Given the determinate development of their nervous systems, 30-some years ago it was taken as given that C. elegans are too simple to learn. However, once researchers turned to examine learning and memory in these tiny animals, they found an incredible amount of flexible behavior and sensitivity to experience. C. elegans have short-term and long-term memory, they can learn through habituation (Rankin et al., 1990), association (Wen et al., 1997), and imprinting (Remy & Hobert, 2005). They pass associative learning tasks using a variety of sensory modalities, including taste, smell, sensitivity to temperature, and sensitivity to oxygen (Ardiel & Rankin, 2010). They also integrate information from different sensory modalities, and respond differently to different levels of intoxicating substances, ā€œsupport[ing] the view that worms can associate a physiological state with a specific experienceā€ (Rankin, 2004, p. R618). There is also behavioral evidence that C. elegans engage in motivational trade-offs. These worms will flexibly choose to head through a noxious environment to gain access to a nutritious substance when hungry enough (Ghosh et al., 2016)—though Birch and colleagues are not convinced this behavior satisfies the marker of motivational trade-offs because it appears that one reflex is merely inhibiting another (Birch et al., 2021, p. 31).

C. elegans are a model organism for the study of nociceptors, and much of what we now know about the mechanisms of nociception comes from studies on this species (Smith & Lewin, 2009). Behavioral responses to noxious stimuli are modulated by opiates, as demonstrated by a study finding that administration of morphine has a dose-dependent effect on the latency of response to heat (Pryor et al., 2007). And, perhaps surprisingly, when the nerve ring that comprises the C. elegans brain was recently mapped, researchers found that different regions of the brain support different circuits that route sensory information to another location where they are integrated, leading to action (Brittin et al., 2021).

Even if we grant the author's low confidence in nematodes' having marker five (motivational trade-offs), current science provides ample confidence that nematodes have markers one (nociceptors), two (integrated brain regions), four (responsiveness to analgesics), and seven (sophisticated associative learning). Given high confidence that nematodes have even three of these markers, the report's methodology would have us conclude that there is ā€œsubstantial evidenceā€ of sentience in nematodes.

The presence of pain markers in C. elegans and in other animals presumed to be unconscious has led to a rejection of some of these markers; for example, Elizabeth Irvine concludes that given the assumption that C. elegans are not conscious, the evidence that they possess nociceptors, engage in motivational trade-offs and show associative learning invalidates these three markers, and raise questions about others (Irvine, 2020). Irvine's worry about a marker's validity would only be strengthened were we to find evidence of it in a brainless animal, such as the sea sponge Porifera, since brainless animals appear to be even worse candidates for sentience. So let us turn to look at the sea sponge.

From the perspective of the lay observer diving through a reef, sponges themselves do nothing. However, sponges reproduce by creating larvae that swim from the parent and later sink and crawl to find a place to settle and metamorphosize into a new sponge. Our current knowledge of sponge larvae behavior is quite slight. What we do know is that the larvae demonstrate negative phototaxis, and their settlement time is increased by the introduction of substrates such as rubble and biofilm into the environment (Wahab et al., 2011). If it were to turn out that sponges engage in motivational trade-offs between light levels and rubble when it comes to selecting a place to settle, would that finding offer more evidence against motivational trade-offs as a marker, should it offer some evidence that sponges are sentient, or should that data imply neither? According to the sentience report's methodology, the answer is neither, but such evidence should not lead us to reject the marker of motivational trade-offs, either. Rather, high confidence in the presence of zero to one markers should lead us toward agnosticism about sentience in the species, and high confidence in one marker coupled with good research that fails to find the other seven markers should lead us to conclude that sentience is unlikely. On the sentience report's approach, and contra Irvine, the markers themselves are not open to being invalidated.

I skimmed the article Disentangling perceptual awareness from nonconscious processing in rhesus monkeys (Macaca mulatta) by Moshe Ben-Haim and others. I liked it, but I believe Kristin's criticism of defining markers of consciousness applies all the same.

I would like the focus to be on comparisons of expected hedonistic welfare across species and substrates (biological or not) instead of their probabilty of sentience. Relatedly, Rethink Priorities (RP) has a research agenda about interspecies welfare comparisons more broadly (not just under expectational total hedonistic utilitarianism).