Octopus bimaculoides genome and transcriptome sequencing demonstrated that a core gene repertoire broadly similar to that of other invertebrate bilaterians is accompanied by expansions in the protocadherin and C2H2 zinc-finger transcription factor families and large-scale genome rearrangements closely associated with octopus-specific transposable elements. Octopuses have been called 'the most intelligent invertebrate', with a host of complex behaviours, and a nervous system comparable in size to that of mammals but organized in a very different manner. It had been hypothesized that, as in vertebrates, whole-genome duplication contributed to the evolution of this complex nervous system. Caroline Albertin et al. have sequenced the genome and multiple transcriptomes of the California two-spot octopus (Octopus bimaculoides) and find no evidence for such duplications but there are large-scale genome rearrangements closely associated with octopus-specific transposable elements. The core developmental and neuronal gene repertoire turns out to be broadly similar to that of other invertebrates, apart from expansions in two gene families formerly thought to be uniquely expanded in vertebrates — the protocadherins (cell-adhesion molecules that regulate neural development) and the C2H2 superfamily of zinc-finger transcription factors. Coleoid cephalopods (octopus, squid and cuttlefish) are active, resourceful predators with a rich behavioural repertoire1. They have the largest nervous systems among the invertebrates2 and present other striking morphological innovations including camera-like eyes, prehensile arms, a highly derived early embryogenesis and a remarkably sophisticated adaptive colouration system1,3. To investigate the molecular bases of cephalopod brain and body innovations, we sequenced the genome and multiple transcriptomes of the California two-spot octopus, Octopus bimaculoides. We found no evidence for hypothesized whole-genome duplications in the octopus lineage4,5,6. The core developmental and neuronal gene repertoire of the octopus is broadly similar to that found across invertebrate bilaterians, except for massive expansions in two gene families previously thought to be uniquely enlarged in vertebrates: the protocadherins, which regulate neuronal development, and the C2H2 superfamily of zinc-finger transcription factors. Extensive messenger RNA editing generates transcript and protein diversity in genes involved in neural excitability, as previously described7, as well as in genes participating in a broad range of other cellular functions. We identified hundreds of cephalopod-specific genes, many of which showed elevated expression levels in such specialized structures as the skin, the suckers and the nervous system. Finally, we found evidence for large-scale genomic rearrangements that are closely associated with transposable element expansions. Our analysis suggests that substantial expansion of a handful of gene families, along with extensive remodelling of genome linkage and repetitive content, played a critical role in the evolution of cephalopod morphological innovations, including their large and complex nervous systems.
The anatomical and biophysical specializations of octopus cells allow them to detect the coincident firing of groups of auditory nerve fibers and to convey the precise timing of that coincidence to their targets. Octopus cells occupy a sharply defined region of the most caudal and dorsal part of the mammalian ventral cochlear nucleus. The dendrites of octopus cells cross the bundle of auditory nerve fibers just proximal to where the fibers leave the ventral and enter the dorsal cochlear nucleus, each octopus cell spanning about one-third of the tonotopic array. Octopus cells are excited by auditory nerve fibers through the activation of rapid, calcium-permeable, alpha-amino-3-hydroxy-5-methyl-4-isoxazole-propionate receptors. Synaptic responses are shaped by the unusual biophysical characteristics of octopus cells. Octopus cells have very low input resistances (about 7 M Omega), and short time constants (about 200 microsec) as a consequence of the activation at rest of a hyperpolarization-activated mixed-cation conductance and a low-threshold, depolarization-activated potassium conductance. The low input resistance causes rapid synaptic currents to generate rapid and small synaptic potentials. Summation of small synaptic potentials from many fibers is required to bring an octopus cell to threshold. Not only does the low input resistance make individual excitatory postsynaptic potentials brief so that they must be generated within 1 msec to sum but also the voltage-sensitive conductances of octopus cells prevent firing if the activation of auditory nerve inputs is not sufficiently synchronous and depolarization is not sufficiently rapid. In vivo in cats, octopus cells can fire rapidly and respond with exceptionally well-timed action potentials to periodic, broadband sounds such as clicks. Thus both the anatomical specializations and the biophysical specializations make octopus cells detectors of the coincident firing of their auditory nerve fiber inputs.
The larvae of the Japanese commercial octopod, Octopus (Octopus) vulgare CUVIER, were successfully reared from the swimming larvae just after hatching to the benthic young octopus in 33 to 40 days on a diet of the zoea larvae of the shrimp, Palaemon serrifer. The rearing experiments were carried out at the Hyogo Prefectural Fisheries Experimental Station from June to September, 1962. At a start of experiments 200 individuals of the octopus larvae were kept in a glass vessel containing about 8 litre of sea-water. During the early 20 days period the sea-water of each vessel was aerated and renewed every 3rd or 4th day, and after this period the method was changed for the use of running water. The results obtained are summarized as follows: 1) From just after hatching, the larvae of the octopus were fed on the larvae of Palaemon of 2 to 4mm body-length. (See Table 1) After transforming into benthic stage, they were given a small piece of ovaries, testes and hepatic glands of the crab, Charybdis japonica. Three or four days after transforming, they were fed on small shrimps and young crabs, Gaetice depressus (DE HAAN) of 5 to 7mm carapace-width. It was observed that the young octopus of 30mm total-length ate 4 or 5 young crabs in a day. 2) The survival rate of the octopus larvae was found as low as 9 percent through the free-swimming stage. However, it would be conceivably possible to raise this survival rate much higher if sufficient food supply is secured during the rearing experiments. 3) The growth of the octopus larvae in 90 days after hatching at water temperature 24.7°C (23.0-26.7°C) can be seen in Table 4 and Fig. 1. It was found that the larvae settled on the bottom 33 to 40 days after hatching, when they attained a size ranging 10-15mm in total-length (3.8-5.7mm in body-length). Once the swimming larvae transformed into the benthic young octopus, they became nocternal and fed on foods mainly at night. A considerable mortality occurred among the young octopus, which was caused by cannibalism and creeping out of the rearing water. 4) The numbers of suckers found on each arm were 3 at the hatching, 5 (4-6) at 10 days, 9 (6-12) at 20 days, 19 (16-22) at 30 days, 24 (21-27) at 40 days and 22-23 when a majority of octopus larvae grew to the benthic stage.
Octopuses of the family Octopodidae adopt two major life-history strategies. The first is the production of relatively few, large eggs resulting in well-developed hatchlings that resemble the adults and rapidly adopt the benthic habit of their parents. The second strategy is production of numerous small eggs that hatch into planktonic, free-swimming hatchlings with few suckers, simple chromatophores and transparent musculature. These distinctive planktonic stages are termed paralarvae and differ from conspecific adults in their morphology, physiology, ecology and behaviour. This study aims to review available knowledge on this subject. In benthic octopuses with planktonic stages, spawning characteristics and duration of planktonic life seem to play an important role in their dispersal capacities. Duration of the hatching period of a single egg mass can range from 2 days to 11 wk, while duration of the planktonic stage can range from 3 wk to half a year, depending on the species and temperature. Thus these paralarvae possess considerable potential for dispersal. In some species, individuals reach relatively large sizes while living as part of the micronekton of oceanic, epipelagic waters. Such forms appear to delay settlement for an unknown period that is suspected to be longer than for paralarvae in more coastal, neritic waters. During the planktonic period, paralarval octopuses feed on crustaceans as their primary prey. In addition to the protein, critical to the protein-based metabolism of octopuses (and all cephalopods), the lipid and copper contents of the prey also appear important in maintaining normal growth. Littoral and oceanic fishes are their main predators and defence behaviours may involve fast swimming speeds, use of ink decoys, dive responses and camouflage. Sensory systems of planktonic stages include photo-, mechano- and chemoreceptors controlled by a highly evolved nervous system that follows the general pattern described for adult cephalopods. On settlement, a major metamorphosis occurs in morphology, physiology and behaviour. Morphological changes associated with the settlement process include positive allometric arm growth; chromatophore, iridophore and leucophore genesis; development of skin sculptural components and a horizontal pupillary response. At the same time, animals lose the Kölliker organs that cover the body surface, the ‘lateral line system’ and the oral denticles of the beaks. Strong positive phototaxis is a common response for hatchlings and some later paralarval stages but this response reduces, disappears or reverses after settlement. There are many gaps in our knowledge of the planktonic phases of benthic octopuses. Most of our understanding of octopus paralarvae comes from studies of just two species (Octopus vulgaris and Enteroctopus dofleini) and knowledge of the vast majority of benthic octopus species with planktonic stages is considered rudimentary or non-existent. Research is needed in a variety of fields, from taxonomy to ecology. Studies of feeding and nutrition are critical in order to develop the nascent aquaculture of key species and ageing studies are necessary to understand planktonic population dynamics,particularly in commercially valuable species targeted by fisheries. Current and potential anthropogenic impacts on these early life stages of octopuses, such as pollution, overfishing and global warming, are also identified.
Octopuses can whip their soft arms with a characteristic "bend propagation" motion to capture prey with sensitive suckers. This relatively simple strategy provides models for robotic grasping, controllable with a small number of inputs, and a highly deformable arm with sensing capabilities. Here, we implemented an electronics-integrated soft octopus arm (E-SOAM) capable of reaching, sensing, grasping, and interacting in a large domain. On the basis of the biological bend propagation of octopuses, E-SOAM uses a bending-elongation propagation model to move, reach, and grasp in a simple but efficient way. E-SOAM's distal part plays the role of a gripper and can process bending, suction, and temperature sensory information under highly deformed working states by integrating a stretchable, liquid-metal-based electronic circuit that can withstand uniaxial stretching of 710% and biaxial stretching of 270% to autonomously perform tasks in a confined environment. By combining this sensorized distal part with a soft arm, the E-SOAM can perform a reaching-grasping-withdrawing motion across a range up to 1.5 times its original arm length, similar to the biological counterpart. Through a wearable finger glove that produces suction sensations, a human can use just one finger to remotely and interactively control the robot's in-plane and out-of-plane reaching and grasping both in air and underwater. E-SOAM's results not only contribute to our understanding of the function of the motion of an octopus arm but also provide design insights into creating stretchable electronics-integrated bioinspired autonomous systems that can interact with humans and their environments.
The octopus sucker represents a fascinating natural system performing adhesion on different terrains and substrates. Octopuses use suckers to anchor the body to the substrate or to grasp, investigate and manipulate objects, just to mention a few of their functions. Our study focuses on the morphology and adhesion mechanism of suckers in Octopus vulgaris. We use three different techniques (MRI, ultrasonography, and histology) and a 3D reconstruction approach to contribute knowledge on both morphology and functionality of the sucker structure in O. vulgaris. The results of our investigation are two-fold. First, we observe some morphological differences with respect to the octopus species previously studied (i.e., Octopus joubini, Octopus maya, Octopus bimaculoides/bimaculatus and Eledone cirrosa). In particular, in O. vulgaris the acetabular chamber, that is a hollow spherical cavity in other octopuses, shows an ellipsoidal cavity which roof has an important protuberance with surface roughness. Second, based on our findings, we propose a hypothesis on the sucker adhesion mechanism in O. vulgaris. We hypothesize that the process of continuous adhesion is achieved by sealing the orifice between acetabulum and infundibulum portions via the acetabular protuberance. We suggest this to take place while the infundibular part achieves a completely flat shape; and, by sustaining adhesion through preservation of sucker configuration. In vivo ultrasonographic recordings support our proposed adhesion model by showing the sucker in action. Such an underlying physical mechanism offers innovative potential cues for developing bioinspired artificial adhesion systems. Furthermore, we think that it could possibly represent a useful approach in order to investigate any potential difference in the ecology and in the performance of adhesion by different species.
Octopuses are generally characterised by rapid non-asymptotic growth, with high individual variability. However, in situ octopus growth is not well understood. The lack of an ageing method has resulted in the majority of our understanding of octopus growth coming from laboratory studies. Despite not being applicable to cephalopods, Modal Progression Analysis (MPA) of length–frequency data is the most common method for examining in situ octopus growth. Recently, counting growth increments in beaks and vestigial shells, and quantifying lipofuscin in brain tissue, have all shown promise for the ageing octopus. Octopuses generally demonstrate two-phase growth in the laboratory, with physiological changes possibly associated with the switch between an initial rapid exponential phase and a slower power growth phase. Temperature and food ration and quality are key factors influencing the initial growth phase. Temperature, however, does not appear to affect the second phase in any consistent way, perhaps because maturity stage can influence the growth response. There may be basic differences in the mechanisms of octopus muscle growth compared with that of other cephalopods. Furthermore, higher relative maintenance energy expenditure, along with the low energy content of their prey, may account for the relatively slow growth of deep-sea octopuses compared to littoral species.
The octopus is a marine animal whose body has no rigid structures. It has eight arms composed of a peculiar muscular structure, named a muscular hydrostat. The octopus arms provide it with both locomotion and grasping capabilities, thanks to the fact that their stiffness can change over a wide range and can be controlled through combined contractions of the muscles. The muscular hydrostat can better be seen as a modifiable skeleton. Furthermore, the morphology the arms and the mechanical characteristics of their tissues are such that the interaction with the environment (i.e., water) is exploited to simplify control. Thanks to this effective mechanism of embodied intelligence, the octopus can control a very high number of degrees of freedom, with relatively limited computing resources. From these considerations, the octopus emerges as a good model for embodied intelligence and for soft robotics. The prototype of a robot arm has been built based on an artificial muscular hydrostat inspired to the muscular hydrostat of the Octopus vulgaris. The prototype presents the morphology of the biological model and the broad arrangement of longitudinal and transverse muscles. Actuation is obtained with cables (longitudinally) and with shape memory alloy springs (transversally). The robot arm combines contractions and it can show the basic movements of the octopus arm, like elongation, shortening and bending, in water.
Octopuses forage far from temporary home dens to which they return for shelter. Spatial tasks may assess learning. Octopuses (Octopus bimaculoides) were placed in a novel arena, and their movements were tracked for 72 hr. Movements around the arena decreased across time, consistent with exploratory learning. Next, octopuses were given 23 hr to move around an arena; after a 24-hr delay, their memory of a burrow location was tested. Most remembered the location of the open burrow, demonstrating learning in 1 day. Finally, octopuses were trained to locate a single open escape burrow among 6 possible locations. Retention was tested after a week and was immediately followed by reversal training (location rotated 180 degrees ). Octopuses learned the original location of the burrow, remembering it for a week. Path lengths increased significantly after reversal, gradually improving and showing relearning. Octopuses show exploratory behavior, learning, and retention of spatial information.
MOTIVATION: As alpha-helical transmembrane proteins constitute roughly 25% of a typical genome and are vital parts of many essential biological processes, structural knowledge of these proteins is necessary for increasing our understanding of such processes. Because structural knowledge of transmembrane proteins is difficult to attain experimentally, improved methods for prediction of structural features of these proteins are important. RESULTS: OCTOPUS, a new method for predicting transmembrane protein topology is presented and benchmarked using a dataset of 124 sequences with known structures. Using a novel combination of hidden Markov models and artificial neural networks, OCTOPUS predicts the correct topology for 94% of the sequences. In particular, OCTOPUS is the first topology predictor to fully integrate modeling of reentrant/membrane-dipping regions and transmembrane hairpins in the topological grammar. AVAILABILITY: OCTOPUS is available as a web server at http://octopus.cbr.su.se.
Octopuses can employ their tapered arms to catch prey of all shapes and sizes due to their dexterity, flexibility, and gripping power. Intrigued by variability in arm taper angle between different octopus species, we explored the utility of designing soft actuators exhibiting a distinctive conical geometry, compared with more traditional cylindrical forms. We find that these octopus-inspired conical-shaped actuators exhibit a wide range of bending curvatures that can be tuned by simply altering their taper angle and they also demonstrate greater flexibility compared with their cylindrical counterparts. The taper angle and bending curvature are inversely related, whereas taper angle and applied bending force are directly related. To further expand the functionality of our soft actuators, we incorporated vacuum-actuated suckers into the actuators for the production of a fully integrated octopus arm-inspired gripper. Notably, our results reveal that because of their enhanced flexibility, these tapered actuators with suckers have better gripping power than their cylindrical-shaped counterparts and require significantly larger forces to be detached from both flat and curved surfaces. Finally, we show that by choosing appropriate taper angles, our tapered actuators with suckers can grip, move, and place a remarkably wide range of objects with flat, nonplanar, smooth, or rough surfaces, as well as retrieve objects through narrow openings. The results from this study not only provide new design insights into the creation of next-generation soft actuators for gripping a wide range of morphologically diverse objects but also contribute to our understanding of the functional significance of arm taper angle variability across octopus species.
The octopus is an interesting model for the development of soft robotics, due to its high deformability, dexterity and rich behavioural repertoire. To investigate the principles of octopus dexterity, we designed an eight-arm soft robot and evaluated its performance with focused experiments. The OCTOPUS robot presented here is a completely soft robot, which integrates eight arms extending in radial direction and a central body which contains the main processing units. The front arms are mainly used for elongation and grasping, while the others are mainly used for locomotion. The robotic octopus works in water and its buoyancy is close to neutral. The experimental results show that the octopus-inspired robot can walk in water using the same strategy as the animal model, with good performance over different surfaces, including walking through physical constraints. It can grasp objects of different sizes and shapes, thanks to its soft arm materials and conical shape.
Recent studies have shown that coastal and shelf cephalopod populations have increased globally over the last six decades. Although cephalopod landings are dominated by the squid fishery, which represents nearly 80% of the worldwide cephalopod catches, octopuses and cuttlefishes represent ∼10% each. Total reported global production of octopuses over the past three decades indicates a relatively steady increase in catch, almost doubling from 179,042 t in 1980 to 355,239 t in 2014. Octopus fisheries are likely to continue to grow in importance and magnitude as many finfish stocks are either fully or over-exploited. More than twenty described octopus species are harvested from some 90 countries worldwide. The current review describes the major octopus fisheries around the globe, providing an overview of species targeted, ecological and biological features of exploited stocks, catches and the key aspects of management.
This paper reports the rationale and design of a robotic arm, as inspired by an octopus arm. The octopus arm shows peculiar features, such as the ability to bend in all directions, to produce fast elongations, and to vary its stiffness. The octopus achieves these unique motor skills, thanks to its peculiar muscular structure, named muscular hydrostat. Different muscles arranged on orthogonal planes generate an antagonistic action on each other in the muscular hydrostat, which does not change its volume during muscle contractions, and allow bending and elongation of the arm and stiffness variation. By drawing inspiration from natural skills of octopus, and by analysing the geometry and mechanics of the muscular structure of its arm, we propose the design of a robot arm consisting of an artificial muscular hydrostat structure, which is completely soft and compliant, but also able to stiffen. In this paper, we discuss the design criteria of the robotic arm and how this design and the special arrangement of its muscular structure may bring the building of a robotic arm into being, by showing the results obtained by mathematical models and prototypical mock-ups.
Comparative analysis of brain function in invertebrates with sophisticated behaviors, such as the octopus, may advance our understanding of the evolution of the neural processes that mediate complex behaviors. Until the last few years, this approach was infeasible due to the lack of neurophysiological tools for testing the neural circuits mediating learning and memory in the brains of octopus and other cephalopods. Now, for the first time, the adaptation of modern neurophysiological methods to the study of the central nervous system of the octopus allows this avenue of research. The emerging results suggest that a convergent evolutionary process has led to the selection of vertebrate-like neural organization and activity-dependent long-term synaptic plasticity. As octopuses and vertebrates are very remote phylogenetically, this convergence suggests the importance of the shared properties for the mediation of learning and memory.
The octopus arm requires special motor control schemes because it consists almost entirely of muscles and lacks a rigid skeletal support. Here we present a 2D dynamic model of the octopus arm to explore possible strategies of movement control in this muscular hydrostat. The arm is modeled as a multisegment structure, each segment containing longitudinal and transverse muscles and maintaining a constant volume, a prominent feature of muscular hydrostats. The input to the model is the degree of activation of each of its muscles. The model includes the external forces of gravity, buoyancy, and water drag forces (experimentally estimated here). It also includes the internal forces generated by the arm muscles and the forces responsible for maintaining a constant volume. Using this dynamic model to investigate the octopus reaching movement and to explore the mechanisms of bend propagation that characterize this movement, we found the following. 1) A simple command producing a wave of muscle activation moving at a constant velocity is sufficient to replicate the natural reaching movements with similar kinematic features. 2) The biomechanical mechanism that produces the reaching movement is a stiffening wave of muscle contraction that pushes a bend forward along the arm. 3) The perpendicular drag coefficient for an octopus arm is nearly 50 times larger than the tangential drag coefficient. During a reaching movement, only a small portion of the arm is oriented perpendicular to the direction of movement, thus minimizing the drag force.
The combination of exploration, habituation, and exploratory play was investigated in octopuses (Octopus dofleini). Eight octopuses were given 10 trials to investigate a floating pill bottle. Exploration consisted of palpation of the object with the arms. Habituation was noticeable in the 1st trial but was more complex across trials. Two octopuses appeared to show exploratory play. This play consisted of aiming water jets through the flexible funnel, which caused regular transport of the object to and return by the aquarium intake current. In this situation, the amount of the 3 activities appeared to be minimally correlated. The results raise questions about the course of habituation, the definition and the extent of play, and the relation of exploratory play to exploration in complex animals
Peter Godfrey-Smith is a leading philosopher of science. He is also an accomplished scuba diver whose underwater videos of warring octopuses have attracted wide notice. In this book, he brings his parallel careers together to tell a bold new story of how nature became aware of itself. Mammals and birds are widely seen as the smartest creatures on earth. But one other branch of the tree of life has also sprouted higher intelligence: the cephalopods, consisting of the squid, the cuttlefish, and above all the octopus. New research shows that these marvelous creatures display remarkable gifts, with each of their tentacles even capable of thinking for itself. What does it mean that higher intelligence on earth has evolved not once but twice? And that the mind of the octopus is nonetheless so different from our own? Combining science and philosophy with first hand accounts of his cephalopod encounters, Godfrey-Smith shows how primitive organisms bobbing in the ocean began sending signals to each other and how these early forms of communication gave rise to the advanced nervous systems that permit cephalopods to change colours and human beings to speak. By tracing the problem of consciousness back to its roots and comparing the human brain to its most alien and perhaps most remarkable animal relative, Godfrey- Smith's Other Minds sheds new light on one of our most abiding mysteries.
Soft robotics is a challenging and promising branch of robotics. It can drive significant improvements across various fields of traditional robotics, and contribute solutions to basic problems such as locomotion and manipulation in unstructured environments. A challenging task for soft robotics is to build and control soft robots able to exert effective forces. In recent years, biology has inspired several solutions to such complex problems. This study aims at investigating the smart solution that the Octopus vulgaris adopts to perform a crawling movement, with the same limbs used for grasping and manipulation. An ad hoc robot was designed and built taking as a reference a biological hypothesis on crawling. A silicone arm with cables embedded to replicate the functionality of the arm muscles of the octopus was built. This novel arm is capable of pushing-based locomotion and object grasping, mimicking the movements that octopuses adopt when crawling. The results support the biological observations and clearly show a suitable way to build a more complex soft robot that, with minimum control, can perform diverse tasks.
Soft robotics is a current focus in robotics research because of the expected capability of soft robots to better interact with real-world environments. As a point of inspiration in the development of innovative technologies in soft robotics, octopuses are particularly interesting 'animal models'. Octopus arms have unique biomechanical capabilities that combine significant pliability with the ability to exert a great deal of force, because they lack rigid structures but can change and control their degree of stiffness. The octopus arm motor capability is a result of the peculiar arrangement of its muscles and the properties of its tissues. These special abilities have been investigated by the authors in a specific study dedicated to identifying the key principles underlying these biological functions and deriving engineering requirements for robotics solutions. This paper, which is the second in a two-part series, presents how the identified requirements can be used to create innovative technological solutions, such as soft materials, mechanisms and actuators. Experiments indicate the ability of these proposed solutions to ensure the same performance as in the biological model in terms of compliance, elongation and force. These results represent useful and relevant components of innovative soft-robotic systems and suggest their potential use to create a new generation of highly dexterous, soft-bodied robots.