External Articles - Behavior and Intelligence Experiments/Observations

Recognizing cephalopod boreholes in shells and the northward spread of Octopus vulgaris Cuvier, 1797 (Cephalopoda, Octopodoidea)
Auke-Florian HIEMSTRA 2015 full article.

INTRODUCTION Aristotle was the first to observe octopuses feed on molluscs (see D’Arcy Thompson, 1910), but it was Fujita who discovered in 1916 that a hole was bored in the shell of cultured pearl oysters prior to its owner being eaten; a behaviour independently discovered by Pilson & Taylor (1961) in laboratory tanks. Octopuses are versatile carnivores with a diverse array of prey, ranging from soft bodied to heavily armoured organisms, such as bivalves, gastropods and crustacean (Nixon, 1987). There are different techniques of penetrating a shell to gain the meat inside (Steer & Semmens, 2003). Enteroctopus dofleini (Wülker, 1910), for example, has four techniques of getting into a clam. If possible, they use the easiest way, according to the optimal foraging model, resorting to drilling only when other methods are unsuccessful (Mather & Anderson, 2007). Regardless of their prey size, Octopus will always try the pulling method first (Fiorito & Gherardi, 1999), but if unsuccessful, it changes its tactics and initiates a drilling response (Hartwick et al., 1978). ...

This article is particularly interesting because it notes that drilling may NOT be solely by the radula.
The drilling activities seem to be carried out by another structure within the buccal mass, namely the small conical teeth on the tip of the muscular salivary papilla (Nixon, 1979a), as not one out of ten octopuses drilled again after surgical removal of their salivary papilla (Nixon 1979a; 1979b). Since the discovery of octopus drillings there has been speculation about possible chemical action on the shell (Fujita, 1916; Pilson & Taylor, 1961; Arnold & Arnold, 1969; Wodinsky 1969; 1973). After comparison of the shell surface it was indeed concluded that some chemical dissolution during drilling may occur (Nixon et al., 1980; Ambrose, 1988).
 
Signal Use by Octopuses in Agonistic Interactions
David Scheel, Peter Godfrey-Smith (pgs), Matthew Lawrence (jugglematt) 2016

Summary
Cephalopods show behavioral parallels to birds and mammals despite considerable evolutionary distance [1 and 2]. Many cephalopods produce complex body patterns and visual signals, documented especially in cuttlefish and squid, where they are used both in camouflage and a range of interspecific interactions [1, 3, 4 and 5]. Octopuses, in contrast, are usually seen as solitary and asocial [6 and 7]; their body patterns and color changes have primarily been interpreted as camouflage and anti-predator tactics [8, 9,10, 11 and 12], though the familiar view of the solitary octopus faces a growing list of exceptions. Here, we show by field observation that in a shallow-water octopus, Octopus tetricus, a range of visible displays are produced during agonistic interactions, and these displays correlate with the outcome of those interactions. Interactions in which dark body color by an approaching octopus was matched by similar color in the reacting octopus were more likely to escalate to grappling. Darkness in an approaching octopus met by paler color in the reacting octopus accompanied retreat of the paler octopus. Octopuses also displayed on high ground and stood with spread web and elevated mantle, often producing these behaviors in combinations. This study is the first to document the systematic use of signals during agonistic interactions among octopuses. We show prima facie conformity of our results to an influential model of agonistic signaling [ 13]. These results suggest that interactions have a greater influence on octopus evolution than has been recognized and show the importance of convergent evolution in behavioral traits.
 
Pull or Push? Octopuses Solve a Puzzle Problem
Jonas N. Richter, Binyamin Hochner, Michael J. Kuba 2016 (PLOS One full article)


Abstract
Octopuses have large brains and exhibit complex behaviors, but relatively little is known about their cognitive abilities. Here we present data from a five-level learning and problem-solving experiment. Seven octopuses (Octopus vulgaris) were first trained to open an L shaped container to retrieve food (level 0). After learning the initial task all animals followed the same experimental protocol, first they had to retrieve this L shaped container, presented at the same orientation, through a tight fitting hole in a clear Perspex partition (level 1). This required the octopuses to perform both pull and release or push actions. After reaching criterion the animals advanced to the next stage of the test, which would be a different consistent orientation of the object (level 2) at the start of the trial, an opaque barrier (level 3) or a random orientation of the object (level 4). All octopuses were successful in reaching criterion in all levels of the task. At the onset of each new level the performance of the animals dropped, shown as an increase in working times. However, they adapted quickly so that overall working times were not significantly different between levels. Our findings indicate that octopuses show behavioral flexibility by quickly adapting to a change in a task. This can be compared to tests in other species where subjects had to conduct actions comprised of a set of motor actions that cannot be understood by a simple learning rule alone.
 
Peripheral injury alters schooling behavior in squid, Doryteuthis pealeii
Megumi Oshima,Theodor Di Pauli von Treuheim, Julia Carroll,Roger T Hanlon,Edgar T Walters,Robyn J Crook 2016 (subscription Science Direct)

Abstract
Animals with detectable injuries are at escalated threat of predation. The anti-predation tactic of schooling reduces individual predation risk overall, but it is not known how schooling behavior affects injured animals, or whether risks are reduced equally for injured animals versus other school members. In this laboratory study we examined the effects of minor fin injury on schooling decisions made by squid. Schooling behavior of groups of squid, in which one member was injured, was monitored over 24 hours. Injured squid were more likely to be members of a school shortly after injury (0.5–2 h), but there were no differences compared with sham-injured squid at longer time points (6–24 h). Overall, the presence of an injured conspecific increased the probability that a school would form, irrespective of whether the injured squid was a member of the school. When groups containing one injured squid were exposed to a predator cue, injured squid were more likely to join the school, but their position depended on whether the threat was a proximate visual cue or olfactory cue. We found no evidence that injured squid oriented themselves to conceal their injury from salient threats. Overall we conclude that nociceptive sensitization after injury changes grouping behaviors in ways that are likely to be adaptive.
 
Number sense and state-dependent valuation in cuttlefish
Tsang-I Yang, Chuan-Chin Chiao 2016 (Subscription Proceedings of the Royal Society B)

Editorial summary on Physics.org

Abstract
Identifying the amount of prey available is an important part of an animal's foraging behaviour. The risk-sensitive foraging theory predicts that an organism's foraging decisions with regard to food rewards depending upon its satiation level. However, the precise interaction between optimal risk-tolerance and satiation level remains unclear. In this study, we examined, firstly, whether cuttlefish, with one of the most highly evolved nervous system among the invertebrates, have number sense, and secondly, whether their valuation of food reward is satiation state dependent. When food such as live shrimps is present, without training, cuttlefish turn toward the prey and initiate seizure behaviour. Using this visual attack behaviour as a measure, cuttlefish showed a preference for a larger quantity when faced with two-alternative forced choice tasks (1 versus 2, 2 versus 3, 3 versus 4 and 4 versus 5). However, cuttlefish preferred the small quantity when the choice was between one live and two dead shrimps. More importantly, when the choice was between one large live shrimp and two small live shrimps (a prey size and quantity trade-off), the cuttlefish chose the large single shrimp when they felt hunger, but chose the two smaller prey when they were satiated. These results demonstrate that cuttlefish are capable of number discrimination and that their choice of prey number depends on the quality of the prey and on their appetite state. The findings also suggest that cuttlefish integrate both internal and external information when making a foraging decision and that the cost of obtaining food is inversely correlated with their satiation level, a phenomenon similar to the observation that metabolic state alters economic decision making under risk among humans.

 
Octopus engineering, intentional and inadvertent
Taylor & Francis Online
David Scheel, P. Godfrey-Smith (@pgs), S. Linquist, S. Chancellor, M. Hing & M. Lawrence

We previously published a description of discovery of a site where octopuses live in an unusually dense collection of individual dens near one another in a bed of scallop shells amid a rock outcrop. We believe the shell bed is an extended midden, accumulated over time by individual octopuses returning to their dens with food. Here we consider what aspects of material collection, den maintenance, and aggregation are intentional for the octopuses, versus inadvertent consequences of individual decisions. Collection of prey items, transport of prey to the den, den excavation, and collection and use of non-prey materials at the den appear to be intentional behaviors. The occurrence of many dens in close aggregation appears to be an inadvertent outcome of the availability of food and the risk of predation in the habitat. Popular media reports have described this site as an ‘city’ designed by octopuses, but that is not an accurate description of the site.
 
Amazing Octopus - Most Intelligent Animal - Series Of Tests - Must Watch- Documentary HD 2017
PlanetEarth HD Documentaries
Amazing Documentary about the unbelievable intelligence of these animals, in this video you will see how these creatures can manipulate their environment, and use there high IQ and brain power to solve a number of situations presented to them by scientists, enjoy!!
 
Cephalopods in Action
Vecchione, Michael and Clyde F.E. Roper 1991

Cephalopods observed from submersibles in the western North Atlantic.
Bulletin of Marine Science, 49(1-2):433-445.

ABSTRACT: Records of 158 observations of cephalopods from submersibles, primarily the JOHNSON SEA-LINK, have been compiled through collaboration with several investigators. These observations include 118 videotape sequences, 58 collected specimens, and numerous shipboard photographs of live animals. At least 33 species have been observed to date; a few species have been observed repeatedly and could be good subjects for directed studies. The methods developed for in situ observation and subsequent collection of specimens with little or no damage allow descriptions of behavior, morphology, physiology, and distribution that are not possible with other methods of collecting.
 
Cephalopod brains: an overview of current knowledge to facilitate comparison with vertebrates
Shuichi Shigeno, Paul Andrews, Giovanna Ponte, Graziano Fiorito 2018 (subscription Frontiers in Physiology)

Cephalopod and vertebrate neural-systems are often highlighted as a traditional example of convergent evolution. Their large brains, relative to body size, and complexity of sensory-motor systems and behavioral repertoires offer opportunities for comparative analysis. Despite various attempts, questions on how cephalopod ‘brains’ evolved and to what extent it is possible to identify a vertebrate-equivalence, assuming it exists, remain still unanswered. Here, we summarize recent molecular, anatomical and developmental data to explore certain features in the neural organization of cephalopods and vertebrates to explore to what extent an evolutionary convergence is likely. Furthermore, and based on whole body and brain axes as defined in early-stage embryos using the expression patterns of homeodomain-containing transcription factors and axonal tractography, we describe a critical analysis of cephalopod neural systems exploring similarities with the cerebral cortex, thalamus, basal ganglia, midbrain, cerebellum, hypothalamus, brain stem and spinal cord of vertebrates. Our overall aim is to promote and facilitate further, hypothesis-driven, studies of cephalopod neural systems evolution.
 
Cephalopod Brains: An Overview of Current Knowledge to Facilitate Comparison With Vertebrates
Shuichi Shigeno1, Paul L. R. Andrews, Giovanna Ponte, Graziano Fiorito 2018 (Frontiers in Physiology, Full article)
Cephalopod and vertebrate neural-systems are often highlighted as a traditional example of convergent evolution. Their large brains, relative to body size, and complexity of sensory-motor systems and behavioral repertoires offer opportunities for comparative analysis. Despite various attempts, questions on how cephalopod ‘brains’ evolved and to what extent it is possible to identify a vertebrate-equivalence, assuming it exists, remain unanswered. Here, we summarize recent molecular, anatomical and developmental data to explore certain features in the neural organization of cephalopods and vertebrates to investigate to what extent an evolutionary convergence is likely. Furthermore, and based on whole body and brain axes as defined in early-stage embryos using the expression patterns of homeodomain-containing transcription factors and axonal tractography, we describe a critical analysis of cephalopod neural systems showing similarities to the cerebral cortex, thalamus, basal ganglia, midbrain, cerebellum, hypothalamus, brain stem, and spinal cord of vertebrates. Our overall aim is to promote and facilitate further, hypothesis-driven, studies of cephalopod neural systems evolution.
 
SLC6A4 binding site and acute prosocial effects of (+/-)-3,4-methylendioxymethamphetamine (MDMA) are evolutionarily conserved in Octopus bimaculoides.
Gul Dolen, Eric Edsinger 2018 (pdf available from bioRxiv)

Abstract
Human and octopus lineages are separated by over 500 million years of evolution, and show divergent anatomical patterns of brain organization. Moreover, while humans exhibit highly complex social behaviors, octopuses are thought to be largely asocial and solitary. Despite these differences, growing evidence suggests that ancient neurotransmitter systems are shared across vertebrate and invertebrate species, and in many cases enable overlapping functions. Here we provide evidence that, as in humans, the atypical amphetamine derivative (+/-)-3,4-methylendioxymethamphetamine (MDMA) enhances acute prosocial behaviors in Octopus bimaculoides. This finding is paralleled by the evolutionary conservation of the serotonin transporter (SERT, encoded by the Slc6A4 gene) binding site of MDMA in the O. bimaculoides genome. Taken together, these data provide evidence that the neural mechanisms subserving social behaviors exist in O. bimaculoides, and indicate that the role of serotonergic neurotransmission in regulating social behaviors is evolutionarily conserved.
 
Variability in the “stereotyped” prey capture sequence of male cuttlefish (Sepia officinalis) could relate to personality differences
Francesca Zoratto, Giulia Cordeschi. Giacomo Grignani. Roberto Bonanni. Enrico Alleva, Giuseppe Nascett, iJennifer A. Mather. Claudio Carere 2018 (subscription Animal Cognition via Springer)

Abstract
Studies of animal personality have shown consistent between-individual variation in behaviour in many social and non-social contexts, but hunting behaviour has been overlooked. Prey capture sequences, especially in invertebrates, are supposed to be quite invariant. In cuttlefish, the attack includes three components: attention, positioning, and seizure. The previous studies indicated some variability in these components and we quantified it under the hypothesis that it could relate to personality differences. We, therefore, analysed predation sequences of adult cuttlefish to test their association with personality traits in different contexts. Nineteen subjects were first exposed to an “alert” and a “threat” test and then given a live prey, for 10 days. Predation sequences were scored for components of the attack, locomotor and postural elements, body patterns, and number of successful tentacle ejections (i.e. seizure). PCA analysis of predatory patterns identified three dimensions accounting for 53.1%, 15.9%, and 9.6% of the variance and discriminating individuals based on “speed in catching prey”, “duration of attack behaviour”, and “attention to prey”. Predation rate, success rate, and hunting time were significantly correlated with the first, second, and third PCA factors, respectively. Significant correlations between capture patterns and responsiveness in the alert and threat tests were found, highlighting a consistency of prey capture patterns with measures of personality in other contexts. Personality may permeate even those behaviour patterns that appear relatively invariant.
 
A community of minds Commentary on Mather on Octopus Mind Bennett L. Schwartz Psychology Florida International University 2019 (pdf)
Abstract: Mather (2019) provides an excellent overview of the literature on octopus perception, cognition, memory, and behavior. Anyone interested in cephalopod cognition and brain organization will find her target article informative and interesting. In this commentary, I challenge the idea that an individual organism must have an individual mind. Given the structure of the octopus brain and their complex behavior, one must consider the possibility that an octopus is a community of minds rather than a single mind.
 

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