Transition paragraph 1:

Article 1 by Young (1959) showed how octopuses learn to attack or avoid objects

based on rewards or shocks. It highlighted the role of the vertical lobe in memory

formation, where learning changes the brain pathways to encourage or stop certain

behaviors. Article 2 takes this further by looking at what's happening in the brain at the

cellular level. Hochner et al. (2003) found that octopuses have long-term potentiation (LTP)

in their brains, a process also seen in vertebrates, which helps in strengthening memory.

This shift from behavior to brain mechanisms shows that octopuses use similar methods to

vertebrates for learning and memory, hinting at a shared evolutionary process.

Article 1 Summary:

Purpose: To study how octopuses learn by associating stimuli with rewards or shocks.

Approach: Used visual stimuli paired with food or shock to observe behavioral changes. Results:

Found that the vertical lobe is crucial for forming and retaining memories. Conclusions: The

vertical lobe ensures long-term memory of learned behaviors.

Article 2 Summary:

Purpose: To explore the cellular basis of learning in the octopus’s brain. Approach:

Examined brain slices of the vertical lobe for synaptic changes. Results: Discovered that

octopuses show long-term potentiation (LTP), like vertebrates. Conclusions: Suggests that

octopuses and vertebrates may have evolved similar learning mechanisms.

Article 2 found that octopuses have a complex brain process called LTP, which supports

learning and memory like vertebrates. Article 3 then explores a bigger idea: whether

octopuses might have some form of consciousness. Ponte et al. (2022) discuss how

octopuses' advanced learning abilities, behaviors, and complex brains might mean they

have a basic awareness or consciousness. This connects the detailed brain processes from

Article 2 to broader functions like perception and decision-making, suggesting that

octopuses’ neural abilities might lead to more complex cognitive functions.

Article 3 Summary:

Purpose: To investigate the possibility of consciousness in cephalopods. Approach: Reviewed

their behaviors, brain structure, and learning abilities. Results: Suggested that cephalopods'

complex brains might support basic forms of consciousness. Conclusions: Cephalopods,

especially octopuses, could have a form of consciousness due to their advanced neural structures.

Article 1 abstract:

An index of the changes in responsiveness of octopuses to moving figures can be obtained by

studying a set of animals and recording the number of them that come out to attack on each

occasion, and the delay. During the period of 1 to 2 h after feeding, untrained octopuses show an

increased tendency to attack. This period of increased attacks corresponds to that for which food

remains within the crop. However, such attacks do not result in the setting up in the memory of

representations that promote later attacks. If food is given after showing a figure the tendency to

attack is greatly increased. It remains at a high level for longer than when the food is given

before showing, but ultimately declines. However, if the figure is presented again with food
within 10 to 24 h the tendency to attack is raised still higher and declines more slowly. After a

number of such presentations the animals come out regularly to attack the figure. The changes in

tendency to respond after each showing of the figure with food thus provide an estimate of the

time course of decay of the increased excitability in certain pathways in the nervous system. The

half-life of the change after the first rewarded presentation is I to 2 h. Similarly, it is possible to

map the time course of the reduced tendency to attack that follows the giving of a shock after

attack on a moving figure. This time depends on the extent to which the figure had previously

been made 'positive' by association with food. By successive feeding and shocking the tendency

to attack a given figure by a group of octopuses can thus be raised and lowered. The effects are

partly general to all moving figures but are greater for those that resemble the figure shown when

the food (or shock) was administered. Using this effect animals can be trained without shocks to

attack certain figures but not others, pathways 'representing' these figures having been

specifically facilitated by feeding. In octopuses without vertical lobes the effects of food or shock

in increasing or lowering the tendency to attack are like those in normal animals but persist for a

shorter time. In these animals lasting representations ensuring or preventing attack are less

readily set up. When food is given as a reward for attacks on one figure, shocks for a different

one, normal octopuses learn within a few trials to attack the one and avoid the other. In animals

without vertical lobes the effect of food is to increase the tendency to attack both figures and the

effect of shocks is to depress the attacks on both. With alternate trials the tendency to attack thus

swings up and down, the animals responding according to the influence of the immediately

previous stimulus. The effect of the vertical lobe is therefore to ensure persistence in the memory

of appropriate distinct representations of the figures and the associated food or shock, so that

there are 'correct' responses after a few trials.

Cellular mechanisms underlying learning and memory were investigated in the octopus using a

brain slice preparation of the vertical lobe, an area of the octopus brain involved in learning and

memory. Field potential recordings revealed long-term potentiation (LTP) of glutamatergic

synaptic field potentials similar to that in vertebrates. These findings suggest that convergent

evolution has led to the selection of similar activity-dependent synaptic processes that mediate

complex forms of learning and memory in vertebrates and invertebrates.

Article 3 abstract:

It is only in recent decades that subjective experience - or consciousness - has become a

legitimate object of scientific inquiry. As such, it represents perhaps the greatest challenge facing

neuroscience today. Subsumed within this challenge is the study of subjective experience in non-

human animals: a particularly difficult endeavor that becomes even more so, as one crosses the

great evolutionary divide between vertebrate and invertebrate phyla. Here, we explore the

possibility of consciousness in one group of invertebrates: cephalopod molluscs. We believe such

a review is timely, particularly considering cephalopods’ impressive learning and memory

abilities, rich behavioral repertoire, and the relative complexity of their nervous systems and

sensory capabilities. Indeed, in some cephalopods, these abilities are so sophisticated that they

are comparable to those of some higher vertebrates. Following the criteria and framework

outlined for the identification of hallmarks of consciousness in non-mammalian species, here we

propose that cephalopods - particularly the octopus - provide a unique test case among

invertebrates for examining the properties and conditions that, at the very least, afford a basal

faculty of consciousness. These include, among others: (i) discriminatory and anticipatory

behaviors indicating a strong link between perception and memory recall; (ii) the presence of
neural substrates representing functional analogs of thalamus and cortex; (iii) the

neurophysiological dynamics resembling the functional signatures of conscious states in

mammals. We highlight the current lack of evidence as well as potentially informative areas that

warrant further investigation to support the view expressed here. Finally, we identify future

research directions for the study of consciousness in these tantalizing animals.

Citations

1. Hochner, B., Brown, E. R., Langella, M., Shomrat, T., & Fiorito, G. (2003). A

Learning and Memory Area in the Octopus Brain Manifests a Vertebrate-Like

Long-Term Potentiation. Journal of Neurophysiology, 90(5), 3547–3554.

https://doi.org/10.1152/jn.00645.2003

2. Ponte, G., Chiandetti, C., Edelman, D. B., Imperadore, P., Pieroni, E. M., & Fiorito, G. (2022).

Cephalopod Behavior: From Neural Plasticity to Consciousness. Frontiers in Systems

Neuroscience, 15. https://doi.org/10.3389/fnsys.2021.787139

3. Young, J. Z. (1960). Unit Processes in the Formation of Representations in the Memory

of Octopus. Proceedings of the Royal Society of London. Series B, Biological

Sciences, 153(950), 1–17