Title: Memory to Consciousness in Cephalopods

Name: Moe Eid

Due Date: 21 October 2024

TA Name: Savannah Kozole

Course: Comparative Physiology

Word Count: 2125

Introduction:

Cephalopods, especially octopuses, are known for their remarkable behavioral

capabilities, including advanced learning, problem-solving, and adaptability. These abilities have

sparked interest in understanding the neural mechanisms that enable such complex behavior,

which could provide insight into the evolution of cognitive functions across species. Octopuses,

as invertebrates with highly developed brains, offer a unique opportunity to explore how

sophisticated neural processes, such as memory formation and decision-making, might have

evolved in parallel with those found in vertebrates. Beyond their intriguing learning and memory

abilities, some researchers have even suggested that cephalopods may possess a form of

consciousness, an idea that challenges long-standing views of invertebrate cognition.

This paper examines the progression of research on cephalopod cognition, focusing on

three key studies that explore different facets of their learning, memory, and potential

consciousness. First, the influential work by Young (1960) explored how octopuses form and

retain memory, focusing on the role of the vertical lobe in this process. Next, Hochner et al.

(2003) investigated the cellular basis of memory in the octopus’s brain, discovering long-term
potentiation (LTP), a process crucial for memory formation in vertebrates. Finally, Ponte et al.

(2022) extended these findings by considering whether the advanced learning abilities and neural

plasticity observed in octopuses might suggest a form of consciousness. This paper follows the

progression from basic learning mechanisms to the possibility of higher-order cognitive

processes, demonstrating how our understanding of cephalopod cognition has evolved over time.

Article 1:

Summary of "Unit Processes in the Formation of Representations in the Memory of Octopus" by

J.Z. Young (1960)

In the paper Unit Processes in the Formation of Representations in the Memory of

Octopus, Young (1960) explores how octopuses form and retain memories, with a specific focus

on their responses to visual stimuli. The central question addressed was how these animals learn

to attack or avoid moving objects based on their past experiences with rewards (food) or

punishments (shock). Young hypothesized that these behaviors are linked to the formation of

memory traces in the nervous system, which would gradually fade unless reinforced. A key

aspect of this study was to determine the role of the vertical lobe, a brain structure thought to be

involved in memory retention, in sustaining these learned behaviors over time.

To test this hypothesis, Young designed behavioral experiments where octopuses were

repeatedly shown moving figures. In some trials, the figures were paired with food rewards,

while in others, an electric shock followed the animals' attack on the figures. The frequency of

attacks and the delay between stimulus presentation and response were recorded to assess how

memories of the stimuli developed and decayed. Additionally, in some experiments, the vertical

lobes of certain animals were surgically removed to determine how this structure impacted
memory retention. By comparing the behavior of normal octopuses with those that had

undergone surgery, Young sought to understand how long-lasting memories were formed and

how this brain structure influenced behavior.

The results demonstrated that octopuses could indeed learn to associate specific visual

stimuli with positive or negative consequences, which influenced their behavior in future trials.

When a figure was paired with food, the octopuses showed an increased likelihood of attacking

it, as indicated by higher attack frequencies and shorter response times compared to untrained

animals. This tendency persisted for about 1 to 2 hours after the initial pairing, but with repeated

exposure, the memory became stronger, leading the octopuses to continue attacking the figure up

to 24 hours later. On the other hand, when an electric shock followed an attack, the octopuses

became less likely to attack that figure again. The strength and persistence of this avoidance

response varied depending on how strongly the figure had been associated with food before the

shock. Over time, if the shock was not repeated, the avoidance behavior diminished, similar to

the decay of positive associations. Interestingly, octopuses that had their vertical lobes removed

exhibited impaired memory retention. These animals were less consistent in their behavior,

responding more to recent experiences than forming lasting memories. This suggests that the

vertical lobe plays a crucial role in maintaining long-term memory representations in octopuses.

Overall, Young's findings supported his hypothesis that memory formation in octopuses

relies on repeated reinforcement and gradually fades without it. The study also emphasized the

importance of the vertical lobe in sustaining memory over time. Without this brain structure, the

animals struggled to retain learned behaviors, highlighting the complexity of memory processes

in cephalopods. These results offered important insights into how invertebrate nervous systems

support memory in ways that are surprisingly similar to those found in vertebrates.
Transition Paragraph 1:

Young (1960) demonstrated how octopuses learn to associate moving objects with

rewards or punishments, such as food or shocks. This research emphasized the critical role of the

vertical lobe in memory formation, showing that changes in brain pathways facilitated learning,

allowing octopuses to either attack or avoid objects based on prior experiences. However, while

this study identified the importance of the vertical lobe in maintaining learned behaviors, it left

open the question of what specific neural processes underlie these memory functions.

Specifically, how do these changes in brain pathways occur at the cellular level? Building on this

foundational work, Hochner et al. (2003) took the investigation further by exploring the cellular

mechanisms in the octopus’s brain that contribute to learning and memory. Their study revealed

that the octopus exhibits long-term potentiation (LTP), a process also found in vertebrates, which

strengthens synaptic connections and underlies memory formation. By shifting the focus from

behavioral responses to the actual mechanisms in the brain, this research suggests that octopuses

and vertebrates may share similar methods for learning and memory, hinting at a convergent

evolutionary process.

Article 2:

Summary of "A Learning and Memory Area in the Octopus Brain Manifests a Vertebrate-Like

Long-Term Potentiation" by Hochner et al. (2003)

The purpose of the study by Hochner et al. (2003) was to investigate the cellular

mechanisms of learning and memory in octopuses by exploring whether long-term potentiation

(LTP), a process that strengthens synapses in response to repeated stimulation, also occurs in the

octopus brain. LTP is a key feature of memory formation in vertebrates, so the authors
hypothesized that octopuses might use a similar mechanism in their nervous systems. They

aimed to fill a gap in our understanding of how cephalopod brains function at a cellular level to

support learning and memory.

To investigate this, the authors focused on the vertical lobe of the octopus brain, an area

known for its role in learning and memory. They took brain slices from this region and

stimulated them electrically to measure the resulting synaptic activity. The specific focus was on

glutamatergic synapses, as glutamate is a neurotransmitter involved in LTP in vertebrates. The

researchers used field potential recordings to detect changes in the strength of synaptic responses

over time. By stimulating the brain slices repeatedly, they could observe whether the synapses

would undergo LTP, indicating a strengthening of synaptic transmission.

The results showed that the vertical lobe of the octopus does indeed exhibit LTP, similar

to what is seen in vertebrate brains. After repeated electrical stimulation, the glutamatergic

synapses became stronger, showing that the octopus brain can enhance synaptic connections in a

way that supports long-term memory formation. This synaptic strengthening persisted for an

extended period, further suggesting that octopuses, like vertebrates, use LTP as a mechanism for

learning and memory. Additionally, the authors noted that the molecular components involved in

octopus LTP, such as glutamate receptors, are very similar to those in vertebrates, reinforcing the

idea that similar processes are at work in both groups despite their evolutionary distance.

From these results, the authors concluded that octopuses use a vertebrate-like mechanism

for memory storage, confirming their hypothesis. The discovery of LTP in the octopus vertical

lobe suggests that this part of the brain plays a crucial role in memory-related synaptic plasticity.

This study provided important evidence that LTP is not unique to vertebrates and may have

evolved independently in different lineages, such as cephalopods and vertebrates, through

convergent evolution. The findings offer new insights into the shared principles of brain function

across a wide range of species, showing that even invertebrates like octopuses can have highly

developed memory systems.

Transition Paragraph 2:

Hochner et al. (2003) revealed that octopuses demonstrate long-term potentiation (LTP)

in their brains, a process also seen in vertebrates that strengthens synaptic connections and

enhances memory retention. This finding raises intriguing questions about the cognitive abilities

of octopuses, as it suggests that their learning and memory mechanisms may be more complex

than previously thought. Building on these insights into brain plasticity, Ponte et al. (2022)

explored whether octopuses could possess a form of consciousness. By examining their

advanced behaviors, intricate brain structure, and learning capacities, the study bridged the gap

between cellular-level brain mechanisms and broader cognitive functions like perception and

decision-making. The results suggest that the sophisticated neural systems in cephalopods,

particularly octopuses, may support basic forms of consciousness.

Article 3:

Summary of "Cephalopod Behavior: From Neural Plasticity to Consciousness" by Ponte et al.

(2022)

The study by Ponte et al. (2022) aimed to explore whether cephalopods, particularly

octopuses, could exhibit basic forms of consciousness. The authors investigated whether the

complex behaviors, learning abilities, and brain structures of these animals might indicate the

presence of conscious experiences, a concept traditionally thought to be limited to vertebrates.

Their hypothesis was that octopuses might demonstrate consciousness through behaviors and
neural processes that are similar to those observed in vertebrates. The research addressed a

knowledge gap regarding how invertebrate nervous systems, particularly cephalopods, could

support complex cognition and awareness.

To address this question, the authors conducted a comprehensive review of studies on

cephalopod behavior, brain anatomy, and neural plasticity. They focused on behaviors such as

problem-solving, tool use, and anticipating future events, which indicate higher-level cognitive

processes. In terms of brain anatomy, the authors examined whether cephalopods possess

structures that could be compared to the thalamus and cortex in vertebrates, which are both key

regions involved in conscious thought. Additionally, they looked at neurophysiological evidence,

such as brain activity patterns in cephalopods, to see if these patterns resembled those associated

with conscious awareness in mammals.

The results of the review highlighted several key findings. Cephalopods, especially

octopuses, have shown advanced problem-solving skills, such as using tools, escaping

enclosures, and even opening jars to access food. These behaviors suggest that octopuses are

capable of learning, memory recall, and decision-making, which are processes linked to

conscious awareness in other animals. Furthermore, octopuses have demonstrated the ability to

anticipate future events and act based on past experiences, indicating cognitive flexibility.

From a neurological perspective, the study identified brain regions in cephalopods that function

similarly to those in vertebrates associated with consciousness. For example, the central brain of

cephalopods has circuits that play a similar role to the vertebrate thalamus, processing sensory

information. Additionally, the study highlighted that neural plasticity, which allows for learning

and adaptation, is highly developed in cephalopods, particularly in their vertical and superior
frontal lobes. This plasticity is crucial for learning and memory, traits often linked to conscious

awareness.

The authors also discussed neurophysiological evidence, including brain activity patterns that

resemble the oscillatory rhythms seen in conscious mammals. These rhythmic brain states are

associated with sensory processing and decision-making, further supporting the idea that

cephalopods may experience some form of awareness.

In conclusion, the authors suggested that octopuses might have a basic form of

consciousness. Although their conscious experience is likely less complex than that of humans,

the behavioral and neurological evidence suggests that these animals possess some level of

awareness, memory, and decision-making. The findings support the hypothesis that

consciousness may have evolved independently in cephalopods and vertebrates through

convergent evolution, driven by similar environmental and survival pressures. This study

provides valuable insight into the evolution of cognitive abilities, suggesting that consciousness

is not exclusive to vertebrates and may exist in different forms across various animal species.

Conclusion:

The research on cephalopod learning, memory, and cognition shows significant progress

in understanding these complex processes, starting with Young’s (1960) foundational work on

memory formation and continuing with the cellular discoveries of Hochner et al. (2003) and the

broader cognitive implications suggested by Ponte et al. (2022). These studies collectively show

that octopuses not only have advanced memory systems but could potentially engage in higher-

order cognitive functions, such as consciousness. The identification of vertebrate-like long-term

potentiation (LTP) in octopuses underscores how similar neural mechanisms for learning and
memory can independently evolve in different species. This provides valuable insights into how

complex brain functions have developed across diverse evolutionary paths. Future research

should focus on further exploring the cognitive capabilities of cephalopods, particularly in

understanding how their behavior and brain processes can expand our knowledge of

consciousness across different species.

 Primary 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

 Abrstact 1:

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 raisecL 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 similar to 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 reward for attacks at 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.

Abstract 2:

A learning and memory area in the octopus brain manifests a vertebrate-like long-term

potentiation. J Neurophysiol 90: 3547–3554, 2003. First published August 13, 2003;

10.1152/jn.00645.2003. 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.

Abstract 3:

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.

TP3:

REVISED

Moe Eid

Lab Instructor: Savannah Kozole

Lab Section: 11360

Date: 30 September 2024

Word Count: 520

Article 1

Summary of "Unit Processes in the Formation of Representations in the Memory of Octopus" by

J.Z. Young (1960)

In the paper Unit Processes in the Formation of Representations in the Memory of Octopus,

Young (1960) explores how octopuses form and retain memories, with a specific focus on their

responses to visual stimuli. The central question addressed was how these animals learn to attack

or avoid moving objects based on their past experiences with rewards (food) or punishments

(shock). Young hypothesized that these behaviors are linked to the formation of memory traces in

the nervous system, which would gradually fade unless reinforced. A key aspect of this study

was to determine the role of the vertical lobe, a brain structure thought to be involved in memory

retention, in sustaining these learned behaviors over time.

To test this hypothesis, Young designed behavioral experiments where octopuses were repeatedly

shown moving figures. In some trials, the figures were paired with food rewards, while in others,

an electric shock followed the animals' attack on the figures. The frequency of attacks and the
delay between stimulus presentation and response were recorded to assess how memories of the

stimuli developed and decayed. Additionally, in some experiments, the vertical lobes of certain

animals were surgically removed to determine how this structure impacted memory retention. By

comparing the behavior of normal octopuses with those that had undergone surgery, Young

sought to understand how long-lasting memories were formed and how this brain structure

influenced behavior.

The results demonstrated that octopuses could indeed learn to associate specific visual stimuli

with positive or negative consequences, which influenced their behavior in future trials. When a

figure was paired with food, the octopuses showed an increased likelihood of attacking it, as

indicated by higher attack frequencies and shorter response times compared to untrained animals.

This tendency persisted for about 1 to 2 hours after the initial pairing, but with repeated

exposure, the memory became stronger, leading the octopuses to continue attacking the figure up

to 24 hours later. On the other hand, when an electric shock followed an attack, the octopuses

became less likely to attack that figure again. The strength and persistence of this avoidance

response varied depending on how strongly the figure had been associated with food before the

shock. Over time, if the shock was not repeated, the avoidance behavior diminished, similar to

the decay of positive associations. Interestingly, octopuses that had their vertical lobes removed

exhibited impaired memory retention. These animals were less consistent in their behavior,

responding more to recent experiences than forming lasting memories. This suggests that the

vertical lobe plays a crucial role in maintaining long-term memory representations in octopuses.

Overall, Young's findings supported his hypothesis that memory formation in octopuses relies on

repeated reinforcement and gradually fades without it. The study also emphasized the importance

of the vertical lobe in sustaining memory over time. Without this brain structure, the animals
struggled to retain learned behaviors, highlighting the complexity of memory processes in

cephalopods. These results offered important insights into how invertebrate nervous systems

support memory in ways that are surprisingly similar to those found in vertebrates.

Article 2

Summary of "A Learning and Memory Area in the Octopus Brain Manifests a Vertebrate-Like

Long-Term Potentiation" by Hochner et al. (2003)

The purpose of the study by Hochner et al. (2003) was to investigate the cellular mechanisms of

learning and memory in octopuses by exploring whether long-term potentiation (LTP), a process

that strengthens synapses in response to repeated stimulation, also occurs in the octopus brain.

LTP is a key feature of memory formation in vertebrates, so the authors hypothesized that

octopuses might use a similar mechanism in their nervous systems. They aimed to fill a gap in

our understanding of how cephalopod brains function at a cellular level to support learning and

memory.

To investigate this, the authors focused on the vertical lobe of the octopus brain, an area known

for its role in learning and memory. They took brain slices from this region and stimulated them

electrically to measure the resulting synaptic activity. The specific focus was on glutamatergic

synapses, as glutamate is a neurotransmitter involved in LTP in vertebrates. The researchers used

field potential recordings to detect changes in the strength of synaptic responses over time. By

stimulating the brain slices repeatedly, they could observe whether the synapses would undergo

LTP, indicating a strengthening of synaptic transmission.

The results showed that the vertical lobe of the octopus does indeed exhibit LTP, similar to what

is seen in vertebrate brains. After repeated electrical stimulation, the glutamatergic synapses
became stronger, showing that the octopus brain can enhance synaptic connections in a way that

supports long-term memory formation. This synaptic strengthening persisted for an extended

period, further suggesting that octopuses, like vertebrates, use LTP as a mechanism for learning

and memory. Additionally, the authors noted that the molecular components involved in octopus

LTP, such as glutamate receptors, are very similar to those in vertebrates, reinforcing the idea

that similar processes are at work in both groups despite their evolutionary distance.

From these results, the authors concluded that octopuses use a vertebrate-like mechanism for

memory storage, confirming their hypothesis. The discovery of LTP in the octopus vertical lobe

suggests that this part of the brain plays a crucial role in memory-related synaptic plasticity. This

study provided important evidence that LTP is not unique to vertebrates and may have evolved

independently in different lineages, such as cephalopods and vertebrates, through convergent

evolution. The findings offer new insights into the shared principles of brain function across a

wide range of species, showing that even invertebrates like octopuses can have highly developed

memory systems.

Article 3

Summary of "Cephalopod Behavior: From Neural Plasticity to Consciousness" by Ponte et al.

(2022)

The study by Ponte et al. (2022) aimed to explore whether cephalopods, particularly octopuses,

could exhibit basic forms of consciousness. The authors investigated whether the complex

behaviors, learning abilities, and brain structures of these animals might indicate the presence of

conscious experiences, a concept traditionally thought to be limited to vertebrates. Their

hypothesis was that octopuses might demonstrate consciousness through behaviors and neural
processes that are similar to those observed in vertebrates. The research addressed a knowledge

gap regarding how invertebrate nervous systems, particularly cephalopods, could support

complex cognition and awareness.

To address this question, the authors conducted a comprehensive review of studies on

cephalopod behavior, brain anatomy, and neural plasticity. They focused on behaviors such as

problem-solving, tool use, and anticipating future events, which indicate higher-level cognitive

processes. In terms of brain anatomy, the authors examined whether cephalopods possess

structures that could be compared to the thalamus and cortex in vertebrates, which are both key

regions involved in conscious thought. Additionally, they looked at neurophysiological evidence,

such as brain activity patterns in cephalopods, to see if these patterns resembled those associated

with conscious awareness in mammals.

The results of the review highlighted several key findings. Cephalopods, especially octopuses,

have shown advanced problem-solving skills, such as using tools, escaping enclosures, and even

opening jars to access food. These behaviors suggest that octopuses are capable of learning,

memory recall, and decision-making, which are processes linked to conscious awareness in other

animals. Furthermore, octopuses have demonstrated the ability to anticipate future events and act

based on past experiences, indicating cognitive flexibility.

From a neurological perspective, the study identified brain regions in cephalopods that function

similarly to those in vertebrates associated with consciousness. For example, the central brain of

cephalopods has circuits that play a similar role to the vertebrate thalamus, processing sensory

information. Additionally, the study highlighted that neural plasticity, which allows for learning

and adaptation, is highly developed in cephalopods, particularly in their vertical and superior
frontal lobes. This plasticity is crucial for learning and memory, traits often linked to conscious

awareness.

The authors also discussed neurophysiological evidence, including brain activity patterns that

resemble the oscillatory rhythms seen in conscious mammals. These rhythmic brain states are

associated with sensory processing and decision-making, further supporting the idea that

cephalopods may experience some form of awareness.

In conclusion, the authors suggested that octopuses might have a basic form of consciousness.

Although their conscious experience is likely less complex than that of humans, the behavioral

and neurological evidence suggests that these animals possess some level of awareness, memory,

and decision-making. The findings support the hypothesis that consciousness may have evolved

independently in cephalopods and vertebrates through convergent evolution, driven by similar

environmental and survival pressures. This study provides valuable insight into the evolution of

cognitive abilities, suggesting that consciousness is not exclusive to vertebrates and may exist in

different forms across various animal species.

TP2:

Transition Paragraph 1:

Young (1960) demonstrated how octopuses learn to associate moving objects with rewards or

punishments, such as food or shocks. This research emphasized the critical role of the vertical

lobe in memory formation, showing that changes in brain pathways facilitated learning, allowing

octopuses to either attack or avoid objects based on prior experiences. However, while this study

identified the importance of the vertical lobe in maintaining learned behaviors, it left open the

question of what specific neural processes underlie these memory functions. Specifically, how do
these changes in brain pathways occur at the cellular level? Building on this foundational work,

Hochner et al. (2003) took the investigation further by exploring the cellular mechanisms in the

octopus’s brain that contribute to learning and memory. Their study revealed that the octopus

exhibits long-term potentiation (LTP), a process also found in vertebrates, which strengthens

synaptic connections and underlies memory formation. By shifting the focus from behavioral

responses to the actual mechanisms in the brain, this research suggests that octopuses and

vertebrates may share similar methods for learning and memory, hinting at a convergent

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.

Transition Paragraph 2:

Hochner et al. (2003) revealed that octopuses demonstrate long-term potentiation (LTP) in their

brains, a process also seen in vertebrates that strengthens synaptic connections and enhances

memory retention. This finding raises intriguing questions about the cognitive abilities of
octopuses, as it suggests that their learning and memory mechanisms may be more complex than

previously thought. Building on these insights into brain plasticity, Ponte et al. (2022) explored

whether octopuses could possess a form of consciousness. By examining their advanced

behaviors, intricate brain structure, and learning capacities, the study bridged the gap between

cellular-level brain mechanisms and broader cognitive functions like perception and decision-

making. The results suggest that the sophisticated neural systems in cephalopods, particularly

octopuses, may support basic forms of consciousness.

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.

Article 2 abstract:
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

TP1:

OCTOPUS LIKES TO LEARN

The three articles start from landmark, progression, to latest.

1. Landmark article, “Unit Processes in the Formation of Representations in the Memory

of Octopus.”
2. Progression article, “A Learning and Memory Area in the Octopus Brain Manifests a

Vertebrate-Like Long-Term Potentiation.”

3. Latest article, “Cephalopod Behavior: From Neural Plasticity to Consciousness.”

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