Animal Cognition (2023) 26:1065–1072

https://doi.org/10.1007/s10071-023-01753-6

ORIGINAL PAPER

Novel object recognition in Octopus maya

Fabian Vergara‑Ovalle1 · Fructuoso Ayala‑Guerrero2 · Carlos Rosas3 · Hugo Sánchez‑Castillo1

Received: 11 March 2022 / Revised: 16 January 2023 / Accepted: 6 February 2023 / Published online: 21 February 2023
© The Author(s) 2023

Abstract
The Novel Object Recognition task (NOR) is widely used to study vertebrates' memory. It has been proposed as an
adequate model for studying memory in different taxonomic groups, allowing similar and comparable results. Although in
cephalopods, several research reports could indicate that they recognize objects in their environment, it has not been tested
as an experimental paradigm that allows studying different memory phases. This study shows that two-month-old and
older Octopus maya subjects can differentiate between a new object and a known one, but one-month-old subjects cannot.
Furthermore, we observed that octopuses use vision and tactile exploration of new objects to achieve object recognition,
while familiar objects only need to be explored visually. To our knowledge, this is the first time showing an invertebrate
performing the NOR task similarly to how it is performed in vertebrates. These results establish a guide to studying object
recognition memory in octopuses and the ontological development of that memory.

Keywords Exploration · Memory · Novel object · Octopus · Recognition · Behavior

Introduction therefore, having these cognitive capacities in each one of

them most likely reflects an evolutionary convergence. This
Cephalopods show a wide variety of behaviors; these behav- convergence has attracted various research groups' attention
iors manifest cognitive capacities such as learning (Fiorito to understanding how cephalopods achieve these capaci-
and Scotto 1992; Tomita and Aoki 2014; Bublitz et al. 2017), ties, mainly memory. One of the strategies to study this is
emotions (Kuba et al. 2006), puzzle-solving (Richter et al. by comparing cephalopods and vertebrates (Shigeno et al.
2016) and individual recognition (Tricarico et al. 2011). 2018). Still, beyond this, the study of memory in cephalo-
Different taxonomic groups share these capacities: insects pods may help us understand the conditions and constraints
(Simons and Tibbetts 2019), crustaceans, arachnids, and ver- required to achieve different types of memory in a nervous
tebrates (Roth 2013). It is important to note that, the nervous system utterly different from that of vertebrates, understand-
system of every one of these groups differs from each other, ing memory as a cognitive capacity that arises from neural
with more than 550 million years of evolutionary history; networks, regardless of its evolutionary history.
To achieve this, it is necessary to establish a task that dif-
ferent taxa can perform, which may be evaluated similarly
* Hugo Sánchez‑Castillo in each of them. A good candidate for this is the novel object
[email protected] recognition (NOR) task (Blaser and Heyser 2015). This task
1
Neuropsychopharmacology Laboratory, Psychology School,
is based on the innate behavior of animals when encounter-
Universidad Nacional Autónoma de México, colonia copilco ing a new object in a familiar environment; the response
universidad, Avenida Universidad 3000, 1er Piso Edif. B. can be exploration or aversion. The NOR task consists of an
Cub B001, Alcaldía de Coyoacan, Ciudad Universitaria, indirect evaluation of memory through discrimination based
Coyoacán, CP 04510 Mexico City, Mexico
on familiarization. Also, this task has several characteristics
2
Neurosciences Laboratory, Psychology School, Universidad that make it suitable for studying the neurobiology of mem-
Nacional Autónoma de México, Ciudad Universitaria,
Coyoacán, 04510 Mexico City, Mexico
ory; for example, it does not require conditioning, has high
3
ecological validity, is performed quickly, it can be divided
Applied Ecophysiology Laboratory, Facultad de Ciencias,
Unidad Multidisciplinaria para Investigación y Educación,
into phases (acquisition, consolidation, and retrieval), as well
Universidad Nacional Autónoma de México, Puerto de as temporality (long and short-term memory) and finally it
abrigo s/n Sisal, Yucatán, Mexico

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1066 Animal Cognition (2023) 26:1065–1072

has been observed that it is an ability shared by all groups of the adult group consisted of captured subjects. The animals
vertebrates and some invertebrates could share it; However, were kept in 125 L tanks (50 cm length, 50 cm width, 50 cm
it still needs to be confirmed experimentally (Blaser and height). Juvenile and adult groups were housed individually,
Heyser 2015). This task has been widely used in vertebrates, and babies were kept all five together in the same housing
for example, in fish such as sharks and zebrafish (May et al. tank.
2016; Toms and Echeverria 2014; Fuss et al. 2014); birds
such as pigeons (Spetch et al. 2006), crows (Stöwe et al. Animal maintenance
2006) and mammals such as Rhesus monkeys (Rajalingham
et al. 2015), mice (Leger et al. 2013) and rats (Mathiasen Octopuses were maintained in artificial seawater (salinity
and DiCamillo 2010). Conversely, NOR has been poorly 3.5%, pH 8, Nitrite 0, Nitrate 25, Ammonia 0, O ­ 2 > 95%)
studied in invertebrates, some similar tests have been car- on a 12–12 h light cycle with white and red led lights. The
ried out in cuttlefishes and insects, like object discrimination experimental tanks or arenas used during the task differ
and context discrimination (Billard et al. 2020; Solvi et al. between groups due to the significant difference in size
2020; Kelman et al. 2008), but not in the same way as the that the octopuses present during their different stages of
NOR task in vertebrates, so there is still much research to development. The group of babies carried out the task in
be done in the field. 2 L (L) opaque plastic containers, (20 cm length, 10 cm
Although in octopuses, the possibility of remembering width, 10 cm height); for the juvenile group, 60 L (40 cm
objects or places has been anecdotally and experimentally length, 40 cm width, 38 cm height) glass containers were
described (Mather 1991), so far, experimental tasks like used, covered by opaque plastic on three sides; finally, for
the NOR task have yet to be standardized. Beyond that, the adult group, 150 L (60 cm length, 50 cm width, 50 cm
octopuses spend weeks or months in the same den and feed height) glass tanks were used, covered by opaque plastic
by foraging. Hence, it would be reasonable to hypothesize on three sides. To avoid the subjects' stress during the
that octopuses can discriminate between a familiar and whole experiment, a den was placed in each housing and
a novel object. That is why in this study, our aim was to experimental tank. The den was placed on the side of the
standardize this task in Octopus maya (Voss and Solis tank, centered. This den must guard the animal against the
1966) at different ages, to obtain a method for the complete light and be of a suitable size so that the entire octopus can
evaluation of novel object recognition memory in this fit inside, and all eight arms can always be touching the
octopus species and at the same time offer a standardized internal surface of the den. The use of this den increases the
task that allows applying NOR task in other cephalopod willingness of the animal s to explore the objects that will
species. be kept for them in the following phases.

Novel object recognition task

Materials and methods
The NOR task was adapted from other animal models
Ethical statement (Lupetow 2017; Ennaceur 2018; Rossato et al. 2019), with
modifications according to the octopus’s behavior.
All the experiments and handling were carried out under The task consisted of three phases: habituation,
the approval of the bioethics committee of the Faculty of familiarization, and test. The three phases were conducted
Psychology, UNAM, and in accordance with the directive consecutively inside the experimental tank. During the
2010/63/EU (European Parliament), considering the habituation phase, the specimen was placed into the arena
recommendations by Fiorito et al. 2015. and allowed to get used to the environment for 24 h.
Previously, our laboratory group has standardized that O.
Subjects maya spends more time resting in its den than crawling
or climbing when familiar with the environment (data not
The subjects were obtained from the Applied Ecophysiology shown). For this reason, it was established that spending
Laboratory of the UMDI, SISAL. O. maya subjects of three more than two continuous hours in its den during the 24 h
different ages were used; a group of five subjects four weeks of habituation would be a criterion to continue to the next
post-hatching (“babies”), another group of five subjects four phase.
months old (“juvenile”), and, the last group of five subjects The familiarization phase began at the end of the
of unknown age but weighing more than 800 g, for which habituation phase. During this phase, two identical objects
were animals in the reproductive stage (“adults”). The were presented; the objects were approximately the size of
babies and juvenile subjects were raised and maintained in the mantle of the subject. Because we observed that a bigger
laboratory conditions until the end of the experiments, while object would provoke aversion, and a smaller object could

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Animal Cognition (2023) 26:1065–1072 1067

be ignored. Since the mantle length varied considerably for (shrimp) and only those subjects that accepted food at the
each experimental group (babies: 1.1 ± 0.3 cm, juveniles: first attempt, could continue to the next phase. As a result,
7.4 ± 0.6, adults: 24.8 ± 3.2 cm), acrylic figures of different all the subjects took the food on the first feeding attempt.
sizes were used (supplementary Fig. 1). These objects were
placed at the same distance from the den and on opposite Statistical analysis
sides. The objects were presented when the subject was
inside or very close to its den. This phase lasted 30 min. The data were evaluated using the discrimination index
This time was standardized to increase the chance that the ((Novel-Familiar)/Total exploration) and the non-parametric
subjects would explore each of the two objects for at least Mann–Whitney U test to compare the exploration between
20 s, as suggested for other taxa (Ennaceur 2018; Rossato groups and Wilcoxon signed-rank test with the software
et al. 2019). STATISTICA 10 to compare the exploration time of the
The testing phase started at the end of the familiarization familiar object with the novel one, considering p < 0.05 as
phase. During this phase, one of the objects was presented statistically significant.
the day before, and a new one with different characteristics
of shape and color was given. Familiar and new objects iden-
tities were counterbalanced between individuals. Opaque Results
and transparent objects were used, with blue, red, and white
colors. No difference was found in test results, regardless of When juveniles or adults, O. maya differentially
the colors used. This phase lasted 30 min, regardless of how explores a known and new object
long they explored each object. During the familiarization
and test phases, the total exploration time and the visual The juvenile and adult groups showed similar behavior
and tactile exploration of the objects were quantified. The with two types of object exploration; visual and tactile
visual exploration consisted of indirectly approaching one guided exploration. After the objects were explored, the
of the objects, with climbs or jumps, and with at least one of animals returned to their dens in a resting position (Sup-
the eyes directed toward the object. These movements can plementary Fig. 2). In the juvenile and adult groups, it was
continue until they get as close to the object as possible but observed that the exploration times were similar between
without touching it. The tactile exploration phase is when both objects during familiarization (Mann–Whitney U test, p
the specimen touches the object with one or more arms, value = 0.28, Z = 1.088). Although the time dedicated to each
considering that these objects were not attached to the tank type of exploration could vary, the five subjects presented an
walls; hence, the subjects could lift or push them. (Fig. 1). approach and manipulation of both objects (Fig. 2A). After
Due to the test duration, the subjects were fed at the 24 h, the test phase was carried out, and it was observed that
end of each day as part of their maintenance in captivity. the adult subjects explored the familiar object for less time,
In addition, octopuses with high levels of stress tend to compared to the novel object (Wilcoxon, p value = .043,
reject food (Fiorito et al. 2015); therefore, as a way of Z: 2.023) (Fig. 2B), while the juvenile explored the novel
corroborating that the animals were not stressed and could object in a tactile way for a significantly less time (Wil-
continue with the test, at the end of each phase they were fed coxon, p value = 0.028, Z: 2.2), but they showed the same

Fig. 1  Experimental diagram of the novel object recognition task for O. maya. During familiarization, the stage of memory acquisition occurs,
while in the test phase, the evocation of memory is observed

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1068 Animal Cognition (2023) 26:1065–1072

Fig. 2  Results of the NOR test in babies (A–C), juveniles (D–F), familiar object (Wilcoxon, p value = 0.043, Z: 2.02 and Wilcoxon,
and adults (G–I) of O. maya. During the familiarization phase (A, D, p value = 0.028, Z: 2.2; respectively). The insets on each test graph
and G), there are no differences between the total exploration time show the discrimination ratio value (DR) = (Time exploring the novel
of both objects. However, it is observed that babies explored for less object – Time exploring the familiar object) / Time of total explora-
than 60 s, while the juvenile and adult groups explored for more than tion). A DR close to 1 is considered a good discrimination between
100 s. In the total exploration during the test phase (B, E, H), only the familiar object and the novel one, while a DR ≤ of 0.5 is consid-
adults (H) show a significant difference (Wilcoxon, p value = 0.041, ered an inability to distinguish between the familiar object and the
Z: 2.03). If only tactile exploration is considered, both juvenile and novel one (Ennanceur 2018; Sivakumaran et al. 2018)
adult groups showed a differential behavior between the novel and

visual exploration time (Wilcoxon, p value = 0.23, Z: 0.94). At four weeks old, O. maya does not behave
Interestingly, it was observed that there was no manipula- differentially concerning the novel object
tion when exploring the familiar object; when the object is and the familiar one
known, only visual exploration is used for its identification,
but not tactile exploration (Fig. 2C). When conducting the baby group test, an immediate “attack”
behavior toward the objects was observed, meaning the
subjects pounced on them immediately as they entered the
water. This reflected in the decrease of start latency during

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Animal Cognition (2023) 26:1065–1072 1069

the test phase (Fig. 3A). This immediate attack behavior tactile. We observed that when the object becomes familiar,
seems to occur regardless of the size of the objects, since the it is practically not explored tactilely. The results suggest
subjects presented it toward the familiar and novel objects that the familiar object can be recognized visually without
but also toward a net (10 cm), a sponge (7 cm) and even the the need to explore it with another sensorial sense. On the
hand of the experimenter during the cleaning of the tank. other hand, when the object is novel, the two most prominent
For this reason, there was no division regarding the explo- senses of the species are used to explore it.
ration phases; only the time in tactile exploration and the Unlike what was observed in the juvenile and adult
total exploration time (the time spent outside the den or in groups, most of the animals in the babies’ group did not
an activity other than the rest) were evaluated. In these sub- show adequate recognition of a novel object. Instead, they
jects, it was observed that although there is an increase in showed an immediate attack behavior toward both objects.
the time they spend in the tactile exploration of the novel We propose that this immediate attack behavior could
object, compared to the familiar object, this difference did have an ecological explanation since getting food can be
not occur in all individuals and is not statistically significant complicated in the octopus’s early stages of life. In such a
(Wilcoxon, p value = 0.068, Z: 1.82) (Fig. 2E). On the other way that the priority is to attack and try to eat anything that
hand, they spent significantly less time exploring than the comes nearby them. On the other hand, juvenile and adult
adult group (Fig. 3B). subjects could prioritize protecting themselves from possible
predators rather than searching for food; therefore, they
achieve a better inhibition of the attack behavior, allowing
Discussion them to take the time to discriminate novel objects from
familiar ones. Also, we hypothesize that at the babies stage,
The novel object recognition task is a simple and short- tactile exploration could be more important, rather than
term task that has been proposed as ideal for studying and visual exploration because of the behavior of individuals to
comparing the memory of different groups of animals. pounce on objects with all eight arms, in a similar way to
However, it must be adapted to the biology of each group the attack behavior reported for other species of octopuses,
(Rosas et al. 2014), considering behavior, the time between for example, O. vulgaris (Shomrat et al. 2008; Zarrella et al.
phases, the characteristics of the objects, and the arena. The 2015).
present study observed that juvenile and adult subjects of These results suggest that O. maya goes through a
O. maya showed novel object recognition. This recognition maturation stage, during which more complex responses are
is observed as a decrease in the total exploration time of the acquired, allowing better discrimination with the objects to
familiar object. The exploration can, at least, be visual and be explored. This should be reflected in the post-hatching

Fig. 3  Latency to explore (A) and total exploration time during the time than the adults (Wilcoxon, p value = 0.021, Z: 2.3); however,
NOR test in “adult” and “baby” groups (B). The whole explora- they did not show differences between the familiarization and test
tion time in adults was shorter during the test phase (Wilcoxon, p phases
value = 0.043, Z: 2.02). The babies group explored significantly less

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1070 Animal Cognition (2023) 26:1065–1072

nervous system maturation of the species (Vergara-Ovalle habituation. Though this is an important difference between
et al. 2022). Something similar was found by Anderson the task in the murine model and octopuses, it does not
et al. 2004 during NOR in rats. As in our work, they also interfere with the NOR task since it is part of the context
used three groups of different ages: weanling, juveniles, throughout the test. Regarding the exploration time, the
and adults. They observed that juveniles and adult rats can adult subjects total exploration time decreased significantly
discriminate between a novel and a familiar object, 24 h during the test phase. Even though in the murine model,
after familiarization, whereas weanling rats could not. This this would normally be considered as a problem of lack of
seems to indicate a process of nervous system maturation, motivation or motor skills during the test phase, in O. maya,
suggesting a similarity between O. maya and rats. It is it can be explained considering that the subjects returned to
important to mention that to achieve an adequate exploration their resting position in the den once the exploration of the
of the rats of different ages during the task, Anderson et al. objects was completed. During the test phase, the familiar
made the task age appropriate by downsizing the objects and object was not always explored since the total exploration
the arena similarly to how it was done with O. maya in the time was reduced because the animals almost only invested
present work. This ontological difference during the NOR time exploring the novel object.
task seems not to be universal; for example, zebrafish larvae In vertebrates, NOR has been associated with the activity
can discriminate objects from the first days after fertilization of the hippocampus, insular cortex, perirhinal cortex, and
(Bruzzone et al. 2020). Perhaps this difference in the ability medial prefrontal cortex (Tanimizu et al. 2017; Rossato
to discriminate from the earliest stages of life corresponds et al. 2019; Cinalli et al. 2020). Although no invertebrate
to the need to mature a more complex nervous system, has such structures, including octopuses like O. maya, this
considering that zebrafish have 1­ 07 neurons (Friedrich et al. and probably other octopuses species show an evolutive
2013), while rats have an accelerated increase in the number convergence with vertebrates, the ability to remember a
of neurons during the first months after birth and reach familiar object. Therefore, some structures in the central
2 × ­108 neurons (Bandeira et al. 2009). Similarly, octopuses brain of octopuses have been related to memory tasks, and
considerably increase their number of neurons during post- similarity has been sought between these and the structures
hatching development to reach 5 × ­108 neurons (Hochner of the vertebrate brain, particularly its vertical lobe has been
2012). compared with the hippocampus of vertebrates (Shomrat
As far as we know, this is the first time this task has been et al. 2015; Shigeno and Ragsdale 2015; Shigeno et al.
used in a species of octopus, and similar adaptations have 2018). While this comparison helps to understand what
only been made in a few invertebrates such as cuttlefish happens in the octopus brain, from our knowledge of other
(Kelman et al. 2008; Billard 2020) and bumblebees (Solvi more studied groups, it is important to consider a complete
et al. 2020). However, these were not strictly NOR tasks system rather than a similarity for each structure.
but adaptations or another type of discrimination. Here, To integrate the knowledge of how the octopus brain
octopuses were tested with a similar task performed in works, studies were made, dividing the supraesophageal
vertebrates, thus facilitating the comparison between both at region of the octopus brain into two systems; one that
a behavioral and cognitive level. Conversely, the effect that includes the vertical lobe and the superior frontal lobe (VL-
"familiarity of context and objects" has on learning in some SF) responsible for visual memory tasks and another that
species of octopus has been widely described (Fiorito et al. involves the buccal lobe and the inferior frontal lobe (Bu-
1998; Borrelli et al. 2020), however, to our knowledge, it had IF), responsible for somatosensory memory tasks (Wells and
not been standardized in a task that could be compared with Young 1975). This division is useful when evaluating tasks
other groups and measurable as a memory test, until now. such as visual discrimination (Sutherland 1962; Tomita et al.
Also, O. maya presents an innate exploration behavior to 2014) or fear of condition (Shomrat et al. 2008); however, in
novel objects, in a similar way to what happens in murine the results of this work, it is clear that in the NOR task, O.
models (Ennaceur 2018), but different from what happens maya uses visual and tactile exploration. Hence, we propose
in other vertebrates such as Danios, who have an aversion that this task might involve both VL-SF and Bu-IF systems.
to novel objects or neophobia (Fuss et al. 2014). Because This is supported by the histological observations in some
juvenile and adult O. maya subjects tend to move slowly, octopuses’ brain atlas (Jung et al. 2018; Vergara-Ovalle
the duration of the different phases was longer than normally et al. 2022), as there is a neuronal connection between the
used in mice and rats. In this work, the duration of each superior frontal lobe and the inferior frontal lobe. This
phase of 30 min is recommended for O. maya, while it might conform to a physical substrate for the connectivity
varies between 1 and 15 min in mammals. (Antunes and of the possible systems required for O. maya to remember
Biala 2012). Another relevant difference between rats and the familiar object. It is giving rise to propose a series of
octopuses is the presence of a den during the test, which experiments that allows corroboration of whether these
is used to avoid the subjects' stress and achieve correct systems participate and the role of the brain's different

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Animal Cognition (2023) 26:1065–1072 1071

structures during the NOR. It would be interesting to see if Bandeira F, Lent R, Herculano-Houzel S (2009) Changing numbers of
visual exploration is essential for NOR in O. maya since this neuronal and non-neuronal cells underlie postnatal brain growth
in the rat. Proc Natl Acad Sci USA 106(33):14108–14113. https://​
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the objects, it does almost not require tactile exploration to Billard P, Clayton N, Jozet-Alves C (2020) Cuttlefish retrieve whether
recognize them. they smelt or saw a previously encountered item. Sci Rep 10:5413.
The present study gives evidence of evolutionary https://​doi.​org/​10.​1038/​s41598-​020-​62335-x
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convergence between vertebrates and O. maya that allows ing approach to the comparative study of memory. Front Behav
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Essentially, this task can be evaluated almost identically Borrelli L, Chiandetti C, Fiorito G (2020) A standardized bat-
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7717/​peerj.​8890
Supplementary Information The online version contains supplemen- Bublitz A, Weinhold S, Strobel S, Dehnhardt G, Hanke F (2017)
tary material available at https://d​ oi.o​ rg/1​ 0.1​ 007/s​ 10071-0​ 23-0​ 1753-6. Reconsideration of serial visual reversal learning in Octopus
(Octopus vulgaris) from a methodological perspective. Front
Acknowledgements Author´s would like to thank M.C. Claudia Physiol 8:54. https://​doi.​org/​10.​3389/​fphys.​2017.​00054
Caamal-Monsreal and the entire working group at the Applied Eco- Cinalli D, Cohen S, Guthrie K, Stackman R (2020) Object recogni-
physiology Laboratory of the UMDI, Facultad de Ciencias, UNAM, tion memory: distinct yet complementary roles of the mouse
Sisal, Yucatán for providing us with the animals, as well as for her CA1 and perirhinal cortex. Front Mol Neurosci 13:527543.
collaboration and support. M. in C. Ignacio Morales-Salas from the https://​doi.​org/​10.​3389/​fnmol.​2020.​527543
Aquarium of the Facultad de Ciencias, UNAM, for his support in the Ennaceur A (2018) Object novelty recognition memory. In: Ennaceur
care and maintenance of the subjects. And finally, I thank Cynthia Paz- A, de Souza Silva MA (eds) Handbook of behavioral neurosci-
Trejo for her assistance in revising the manuscript. ence. Vol. 27. Handbook of object novelty recognition. Elsevier
Academic Press, pp 1–22
Funding José Fabián Vergara Ovalle would like to thank the program Fiorito G, Scotto P (1992) Observational Learning in Octopus vul-
Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma garis. Science (new York, NY) 256:545–547. https://​doi.​org/​10.​
de México (UNAM) and Consejo Nacional de Ciencia y Tecnología 1126/​scien​ce.​256.​5056.​545
(CONACYT) for the fellowship granted. We thank the projects PAPIIT Fiorito G, Biederman GB, Davey VA, Gherardi F (1998) The role of
IN 306918, IN208722, and PAPIME PE300918 granted to HSC for the stimulus preexposure in problem solving by Octopus vulgaris.
funding provided for this project. Anim Cogn 1(2):107–112. https://​doi.​org/​10.​1007/​s1007​10050​
015
Data availability All data are available from the corresponding authors Fiorito G, Affuso A, Basil J, de Cole A, Girolamo P, D’Angelo L,
upon reasonable request. Dickel L, Gestal C, Grasso F, Kuba M, Mark F, Melillo D,
Osorio D, Perkins K, Ponte G, Shashar N, Smith D, Smith J,
Open Access This article is licensed under a Creative Commons Andrews PL (2015) Guidelines for the Care and Welfare of
Attribution 4.0 International License, which permits use, sharing, Cephalopods in Research -a consensus based on an initiative
adaptation, distribution and reproduction in any medium or format, by CephRes, FELASA and the Boyd Group. Lab Anim 49(2
as long as you give appropriate credit to the original author(s) and the Suppl):1–90. https://​doi.​org/​10.​1177/​00236​77215​580006
source, provide a link to the Creative Commons licence, and indicate Friedrich RW, Genoud C, Wanner AA (2013) Analyzing the structure
if changes were made. The images or other third party material in this and function of neuronal circuits in zebrafish. Front Neural Cir-
article are included in the article's Creative Commons licence, unless cuits 7:71. https://​doi.​org/​10.​3389/​fncir.​2013.​00071
indicated otherwise in a credit line to the material. If material is not Fuss T, Bleckmann H, Schluessel V (2014) Visual discrimination
included in the article's Creative Commons licence and your intended abilities in the gray bamboo shark (Chiloscyllium griseum).
use is not permitted by statutory regulation or exceeds the permitted Zoology (jena) 117:104–111. https://​d oi.​o rg/​1 0.​1 016/j.​z ool.​
use, you will need to obtain permission directly from the copyright 2013.​10.​009
holder. To view a copy of this licence, visit http://​creat​iveco​mmons.​ Hochner B (2012) An embodied view of octopus neurobiology. Curr
org/​licen​ses/​by/4.​0/. Biol 22(20):R887–R892. https://​doi.​org/​10.​1016/j.​cub.​2012.​09.​
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