# Comparative Cognition and the Convergent Evolution of Minds

> Part I: The Architecture of an Alien Mind · Chapter 12 of 17 — The Octopus Mind
> Canonical: https://octopuscognition.org/sections/comparative-cognition-and-the-convergent-evolution-of-minds/

## In brief

Cephalopods are the strongest natural experiment we have in the independent evolution of a mind. The last common ancestor of octopuses and humans lived roughly 550–600 million years ago and was almost certainly a small, flattened, wormlike bilaterian with, at most, a diffuse nerve net—nothing resembling a complex brain (Godfrey-Smith, Other Minds, 2016).

Cephalopods are the strongest natural experiment we have in the independent evolution of a mind. The last common ancestor of octopuses and humans lived roughly 550–600 million years ago and was almost certainly a small, flattened, wormlike bilaterian with, at most, a diffuse nerve net—nothing resembling a complex brain (Godfrey-Smith, *Other Minds*, 2016). This means large brains and sophisticated behavior arose at least twice on Earth, in lineages separated for more than half a billion years: once toward vertebrates (mammals, birds) and once toward coleoid cephalopods. Godfrey-Smith's widely cited framing is that meeting an octopus is "probably the closest we will come to meeting an intelligent alien," and that cephalopod minds are "the most other of all." His book is philosophy and synthesis, not primary data, but it crystallized the comparative-cognition agenda.

**A different architecture reaching similar ends.** An octopus has roughly 500 million neurons—comparable to a dog—but only about a third sit in the central brain; the remaining ~two-thirds are distributed through the eight arms in axial nerve cords and ganglia (often popularized, loosely, as "nine brains"). Arms can execute grasping, chemotactile search, and reaching semi-autonomously, and severed arms continue coordinated behavior, indicating genuinely decentralized control (see Gutnick, Hochner and colleagues; Bilateria arm-atlas work, bioRxiv 2024). This is a fundamentally different way to build cognition than the centralized vertebrate plan, yet it supports jar-opening, maze navigation, observational-style learning, and behavioral flexibility.

**Convergence on specific cognitive capacities.** The most striking evidence is that cephalopods independently evolved capacities long treated as hallmarks of "big-brained" birds and mammals. Finn, Tregenza and Norman (*Current Biology*, 2009) documented *Amphioctopus marginatus* carrying coconut-shell halves to assemble later as shelter—defensive/anticipatory tool use requiring awkward "stilt-walking." Jozet-Alves, Bertin, and—tellingly—Nicola Clayton (*Current Biology*, 2013), the same researcher who established episodic-like ("what-where-when") memory in scrub jays, demonstrated episodic-like memory in cuttlefish, a direct convergence with corvids. Schnell, Hanlon, Clayton et al. (*Proc. R. Soc. B*, 2021) showed common cuttlefish (*Sepia officinalis*) passing a version of the Stanford "marshmallow test," delaying gratification up to ~50–130 seconds—the first invertebrate self-control evidence, and the first non-primate link between self-control and learning performance. Schnell et al. (2021, *Proc. R. Soc. B*) also found episodic-like memory is *preserved* with age in cuttlefish, unlike the age-related decline seen in humans and other mammals—a genuine divergence, not just convergence. The broad comparative case is reviewed in Schnell, Amodio, Boeckle & Clayton, "How intelligent is a cephalopod?" (*Biological Reviews*, 2021).

**Convergence at the level of brain circuits.** J.Z. Young noted decades ago that the octopus vertical lobe is morphologically analogous to the vertebrate hippocampus and insect mushroom body. Shomrat, Hochner and colleagues (2008 and later) showed the vertical lobe uses a vertebrate-like activity-dependent long-term potentiation (LTP) and a "fan-out–fan-in" divergence–convergence connectivity, mirroring associative memory circuits in birds and mammals (reviewed in Hochner & Shomrat). Whether this reflects deep homology or true convergence remains genuinely unresolved.

**Genomic and molecular novelties.** Albertin, Ragsdale et al. (*Nature*, 2015) sequenced *Octopus bimaculoides* and found no whole-genome duplication (the mechanism often invoked for vertebrate complexity); instead a ~168-member protocadherin expansion (~10× other invertebrates, >2× mammals), C2H2 zinc-finger expansion, and massive genome rearrangement. Because cephalopod neurons lack myelin, short-range protocadherin-mediated wiring may have been key. Liscovitch-Brauer, Rosenthal & Eisenberg (*Cell*, 2017) showed coleoids recode >60% of neural transcripts via A-to-I RNA editing (vs <1% in humans), trading genome evolvability for transcriptome plasticity; Birk et al. (*Cell*, 2023) showed temperature-dependent editing dynamically recodes the neural proteome.

**The consciousness debate.** The Cambridge Declaration on Consciousness (Low, Edelman, Koch; 7 July 2012) explicitly named octopuses among animals possessing "neurological substrates that generate consciousness." The broader New York Declaration on Animal Consciousness (NYU, April 2024; Andrews, Birch, Sebo; 500+ signatories) states there is "at least a realistic possibility" of conscious experience in cephalopods and other invertebrates, and that dismissing this possibility is irresponsible for welfare decisions. This scientifically underpinned the UK's inclusion of cephalopods and decapods as sentient in the 2022 Animal Welfare (Sentience) Act (following the Birch LSE review). Debates persist: some (e.g., critics of "agnostic" over-attribution) caution that convergent behavior need not imply subjective experience, and cephalopods' short, largely asocial lives make the standard "social intelligence" and "long-life" drivers of cognition poor fits—an unsolved puzzle Amodio, Clayton, Fiorito et al. framed in "Grow smart and die young" (*Trends in Ecology & Evolution*, 2019).

**Striking / counterintuitive:**
- Octopuses and humans last shared an ancestor ~550-600 Myr ago that had essentially no complex brain—so large brains evolved from near-scratch at least twice, making cephalopods a true independent origin of mind.
- About two-thirds of an octopus's ~500 million neurons are in its arms, not its central brain; severed arms continue coordinated behavior—cognition is genuinely decentralized.
- Cuttlefish episodic-like memory does NOT decline with age (Schnell et al. 2021), the opposite of the memory decline seen in aging humans, mammals, and corvids.
- Coleoid cephalopods recode over 60% of their neural RNA transcripts via A-to-I editing (vs under 1% in humans), and appear to have traded genomic evolvability for this transcriptome plasticity.
- The 2013 cuttlefish episodic-memory paper was co-authored by Nicola Clayton—the very scientist who first demonstrated the same capacity in scrub jays—making the convergence almost poetically direct.
- The octopus genome has ~168 protocadherin genes (roughly 10x other invertebrates, >2x mammals), a gene family previously thought to be a vertebrate specialty for wiring brains.
- Cephalopods break the standard theories of why intelligence evolves: they are mostly short-lived (1-2 years) and asocial, contradicting both the 'social intelligence' and 'long lifespan' hypotheses.

**Open questions:**
- Is the octopus vertical lobe's resemblance to the vertebrate hippocampus/insect mushroom body true convergence, or does it reflect a deep, conserved genetic toolkit (deep homology)?
- Does convergent complex behavior (self-control, episodic-like memory, tool use) actually entail subjective/phenomenal consciousness, or can it arise without felt experience?
- What selective pressures drove cephalopod intelligence given their short, largely solitary lives, which defy the social-brain and long-life hypotheses?
- How is unified behavior (and any unified experience) produced from a radically decentralized nervous system where the arms have substantial autonomy—where, if anywhere, is it 'like something' to be an octopus?
- How much of cephalopod neural plasticity depends on dynamic RNA recoding rather than DNA-encoded circuitry, and what does that imply for comparing their 'intelligence' to genome-based vertebrate cognition?
- Are current cognitive tests (designed for vertebrates) valid measures for such an alien body plan, or do they systematically mis-estimate cephalopod minds?

*Key researchers/labs: Peter Godfrey-Smith (philosopher of biology, USydney/CUNY; Other Minds), Nicola S. Clayton (comparative cognition, Cambridge; episodic-like memory in corvids and cuttlefish), Alexandra K. Schnell (Cambridge/MBL; cuttlefish self-control and comparative cognition), Binyamin Hochner and Tal Shomrat (Hebrew University; vertical lobe, octopus LTP), Graziano Fiorito (Stazione Zoologica Anton Dohrn, Naples; cephalopod learning/behavior), Clifton W. Ragsdale and Caroline B. Albertin (UChicago/MBL; octopus genome), Joshua J.C. Rosenthal and Eli Eisenberg (MBL Woods Hole / Tel Aviv; RNA editing), Roger Hanlon (MBL; cephalopod behavior and camouflage), Jennifer Mather (Lethbridge; octopus personality, play, cognition), Jonathan Birch and Kristin Andrews (LSE / York; animal sentience, NY Declaration), Piero Amodio (comparative cognition; cephalopod intelligence evolution), Christelle Jozet-Alves (Caen; cuttlefish memory).*

### Key papers
- **Albertin CB, Simakov O, Rokhsar DS, Ragsdale CW, et al. (2015).** *The octopus genome and the evolution of cephalopod neural and morphological novelties.* Nature — First cephalopod genome: no whole-genome duplication, but ~168 protocadherins and C2H2 zinc-finger expansions and genome rearrangement underlie neural complexity
- **Liscovitch-Brauer N, Alon S, Rosenthal JJC, Eisenberg E, et al. (2017).** *Trade-off between Transcriptome Plasticity and Genome Evolution in Cephalopods.* Cell — Coleoids recode >60% of neural RNA transcripts via A-to-I editing (vs <1% in humans), trading genomic evolvability for transcriptome plasticity
- **Jozet-Alves C, Bertin M, Clayton NS (2013).** *Evidence of episodic-like memory in cuttlefish.* Current Biology — Cuttlefish show what-where-when memory—direct convergence with corvids, co-authored by the researcher who established it in scrub jays
- **Schnell AK, Boeckle M, Rivera M, Clayton NS, Hanlon RT (2021).** *Cuttlefish exert self-control in a delay of gratification task.* Proceedings of the Royal Society B — First invertebrate evidence of self-control (marshmallow test) and a self-control–learning link outside primates
- **Godfrey-Smith P (2016).** *Other Minds: The Octopus, the Sea, and the Deep Origins of Consciousness.* Farrar, Straus and Giroux — Defining synthesis framing cephalopods as an independent, ~600-Myr-divergent origin of mind—'the closest we will come to meeting an intelligent alien'
- **Finn JK, Tregenza T, Norman MD (2009).** *Defensive tool use in a coconut-carrying octopus.* Current Biology — Documented anticipatory tool use in Amphioctopus marginatus, a benchmark of flexible cognition shared with apes and corvids
- **Shomrat T, Zarrella I, Fiorito G, Hochner B (2008).** *The octopus vertical lobe modulates short-term learning rate and uses LTP to acquire long-term memory.* Current Biology — Shows the vertical lobe uses vertebrate-like LTP, evidence of convergent associative-memory circuitry
- **Amodio P, Boeckle M, Schnell AK, Ostojíc L, Fiorito G, Clayton NS (2019).** *Grow Smart and Die Young: Why Did Cephalopods Evolve Intelligence?.* Trends in Ecology & Evolution — Frames the puzzle that short, asocial cephalopod lives don't fit the standard social/long-life drivers of intelligence
- **Schnell AK, Amodio P, Boeckle M, Clayton NS (2021).** *How intelligent is a cephalopod? Lessons from comparative cognition.* Biological Reviews — Comprehensive review benchmarking cephalopod cognition against vertebrate comparative-cognition standards
- **Birk MA, Liscovitch-Brauer N, Rosenthal JJC, et al. (2023).** *Temperature-dependent RNA editing in octopus extensively recodes the neural proteome.* Cell — Editing dynamically remodels the neural proteome with temperature—a plasticity mechanism absent in vertebrates
- **Low P, Panksepp J, Edelman D, Koch C, et al. (2012).** *The Cambridge Declaration on Consciousness.* Francis Crick Memorial Conference — Formal scientific statement explicitly naming octopuses as possessing neural substrates of consciousness
- **Andrews K, Birch J, Sebo J, et al. (2024).** *The New York Declaration on Animal Consciousness.* NYU (500+ signatories) — Asserts a 'realistic possibility' of consciousness in cephalopods and other invertebrates, with welfare-precautionary implications

## Resolved source links

- [The octopus genome and the evolution of cephalopod neural and morphological novelties.](https://doi.org/10.1038/nature14668) — DOI 10.1038/nature14668
- [Trade-off between Transcriptome Plasticity and Genome Evolution in Cephalopods.](https://doi.org/10.1016/j.cell.2017.03.025) — DOI 10.1016/j.cell.2017.03.025
- [Evidence of episodic-like memory in cuttlefish.](https://doi.org/10.1016/j.cub.2013.10.021) — DOI 10.1016/j.cub.2013.10.021
- [Cuttlefish exert self-control in a delay of gratification task.](https://doi.org/10.1098/rspb.2020.3161) — DOI 10.1098/rspb.2020.3161
- [Other Minds: The Octopus, the Sea, and the Deep Origins of Consciousness.](https://us.macmillan.com/books/9780374537197/otherminds/)
- [Defensive tool use in a coconut-carrying octopus.](https://doi.org/10.1016/j.cub.2009.10.052) — DOI 10.1016/j.cub.2009.10.052
- [The Octopus Vertical Lobe Modulates Short-Term Learning Rate and Uses LTP to Acquire Long-Term Memory.](https://doi.org/10.1016/j.cub.2008.01.056) — DOI 10.1016/j.cub.2008.01.056
- [Grow Smart and Die Young: Why Did Cephalopods Evolve Intelligence?.](https://doi.org/10.1016/j.tree.2018.10.010) — DOI 10.1016/j.tree.2018.10.010
- [How intelligent is a cephalopod? Lessons from comparative cognition.](https://doi.org/10.1111/brv.12651) — DOI 10.1111/brv.12651
- [Temperature-dependent RNA editing in octopus extensively recodes the neural proteome.](https://doi.org/10.1016/j.cell.2023.05.004) — DOI 10.1016/j.cell.2023.05.004
- [The Cambridge Declaration on Consciousness.](https://fcmconference.org/img/CambridgeDeclarationOnConsciousness.pdf)
- [The New York Declaration on Animal Consciousness.](https://sites.google.com/nyu.edu/nydeclaration)

## Related trails

- [The Alien Mind Trail](https://octopuscognition.org/trails/alien-mind/index.md): What does intelligence look like when it is not built like us?