Development of Biomimetic Squid-Inspired Suckers

Biomechanical properties of squid suckers were studied to provide inspiration for the development of sucker artefacts for a robotic octopus.Mechanical support of the rings found inside squid suckers was studied by bending tests.Tensile tests were carried out to study the maximum possible sucking force produced by squid suckers based on the strength of sucker stalks,normalized by the sucking areas.The squid suckers were also directly tested to obtain sucking forces by a special testing arrangement.Inspired by the squid suckers,three types of sucker artefacts were developed for the arm skin of an octopus inspired robot.The first sucker artefact made of knitted nylon sheet reinforced silicone rubber has the same shape as the squid suckers.Like real squid suckers,this type of artefact also has a stalk that is connected to the arm skin and a ring to give radial support.The second design is a straight cylindrical structure with uniform wall thickness made of silicone rubber.One end of the cylinder is directly connected to the arm skin and the other end is open.The final design of the sucker has a cylindrical base and a concave meniscus top.The meniscus was formed naturally using the surface tension of silicone gel,which leads to a higher level of the liquid around the edge of a container.The wall thickness decreases towards the tip of the sucker opening.Sucking forces of all three types of sucker artefacts were measured.Advantages and disadvantages of each sucker type were discussed.The final design of suckers has been implemented to the arm skin prototypes.     

Inspiration, simulation and design for smart robot manipulators from the sucker actuation mechanism of cephalopods

Octopus arms house 200-300 independently controlled suckers that can alternately afford an octopus fine manipulation of small objects and produce high adhesion forces on virtually any non-porous surface. Octopuses use their suckers to grasp, rotate and reposition soft objects (e.g., octopus eggs) without damaging them and to provide strong, reversible adhesion forces to anchor the octopus to hard substrates (e.g., rock) during wave surge. The biological 'design' of the sucker system is understood to be divided anatomically into three functional groups: the infundibulum that produces a surface seal that conforms to arbitrary surface geometry; the acetabulum that generates negative pressures for adhesion; and the extrinsic muscles that allow adhered surfaces to be rotated relative to the arm. The effector underlying these abilities is the muscular hydrostat. Guided by sensory input, the thousands of muscle fibers within the muscular hydrostats of the sucker act in coordination to provide stiffness or force when and where needed. The mechanical malleability of octopus suckers, the interdigitated arrangement of their muscle fibers and the flexible interconnections of its parts make direct studies of their control challenging. We developed a dynamic simulator (ABSAMS) that models the general functioning of muscular hydrostat systems built from assemblies of biologically constrained muscular hydrostat models. We report here on simulation studies of octopus-inspired and artificial suckers implemented in this system. These simulations reproduce aspects of octopus sucker performance and squid tentacle extension. Simulations run with these models using parameters from man-made actuators and materials can serve as tools for designing soft robotic implementations of man-made artificial suckers and soft manipulators.     

The association between distribution and octopodid morphology: implications for classification

The Octopodidae occur in virtually all benthic marine habitats; however, species in the family show little overt morphological differentiation. Subfamilies are currently defined by the presence or absence of an ink sac and the number of sucker rows (the presence of an ink sac and a single row of suckers are primitive characters) (Voss, 1988b); subfamily depth ranges arc cited in the diagnoses. Examination of external octopodid morphology through principal components analysis reveals that octopodid morphology correlates with geographic distribution. Low-latitude, shallow-water octopuses typically have narrower bodies and larger suckers on longer arms than do deep sea and high-latitude species. Sucker size inversely correlates with depth distribution, as studies of sucker functional morphology predict (Kier & Smith, 1990). The same characters contribute in a very similar manner to the discrimination of species when grouped by subfamily and when grouped by mean depth distribution. That depth distributions, which correlate with morphology and with the loss of the ink sac, contribute to the definition of these subfamilies, suggests that the subfamilies constitute phenetically similar rather than monophyletic groups. Cladistic analysis is required to reassess octopodid phylogeny.     

The Morphology and Adhesion Mechanism of Octopus vulgaris Suckers

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.     

Morphological and genetic structuring in the Utah Lake sucker complex

Population decline in the federally endangered June sucker (Chasmistes liorus), a lakesucker unique to Utah Lake, Utah, has been attributed in part to hybridization with the more widespread Utah sucker (Catostomus ardens). As a group, suckers in Utah Lake exhibit considerable external morphological variation. Meristic and morphological ambiguities, presumably the result of hybridization, create a continuum of intermediate forms between Chasmistes and Catostomus extremes and prevent definitive identification to species. Here we describe and evaluate the morphological and genetic variation in suckers in Utah Lake by comparing a morphological analysis with amplified fragment length polymorphism and microsatellite analyses. Suckers were morphologically differentiated using mouth characters associated with different feeding strategies: planktivory (June sucker) and benthivory (Utah sucker). Although we found no genetic evidence for a deep divergence between June and Utah morphs, significant, but slight population structuring accompanied the substantial morphological variation. Bayesian model‐based genetic clustering analyses detected two sucker populations in Utah Lake; however, these clusters were not strongly concordant with morphological groupings or between marker systems. The suckers in Utah Lake present an interesting dilemma regarding conservation: should one conserve (breed and stock) a subset of the morphotypic variation in the Utah Lake sucker complex, focusing on the endangered June sucker morphotype, or should one conserve both June sucker and Utah sucker morphotypes in this complex, possibly maximizing evolutionary potential? We explore this question in the context of current genetic and morphological variation in the Utah Lake sucker complex as well as historical information on this complex and other lakesuckers.     

A NEW SPECIES OF GRANELEDONE (CEPHALOPODA: OCTOPODIDAE) FROM THE SOUTHWEST ATLANTIC OCEAN

A new species of eledonid octopus is described from the southwesternAtlantic Ocean from depths between 90 and 1000 m off the coastsof southern Brazil, Uruguay, and Argentina. This species, Graneledoneyamana is characterized by having a papillose skin, two welldeveloped 'horns' above the eyes, small gills with 5-7 lamellaeon the outer demibranch, arms with uniserial suckers, 35-80on females and 26-70 on males. The third right arm is hectocotylized,the ligula is small, the calamus is large and well differentiated.Ink sac absent. These characters differ from all other knownGraneledone species from the southern oceans. (Received 22 July 1999; accepted 20 May 2000)     

Biology of suckers: late-formed shoots in sugarcane

Suckers in sugarcane are tillers that form late in the growing season after the population of main stalks has been established. Their biology has rarely been studied, though unsubstantiated comments relating to their morphology, development and number are abundant. In this paper, the literature is reviewed and new experiments presented. Comparison of suckers, emerging plant and ratoon cane showed that suckers have a different leaf and stem morphology. Suckers have a greater diameter at their stem base and shorter and wider leaves than primary stalks of a similar age. In 1 ‐year‐old crops, suckers have been demonstrated to decrease the net sucrose concentration of harvested material by diluting sucrose in the main stalks. By comparing main stalks without suckers to those that have initiated suckers, evidence is presented that suckers may also be decreasing the sucrose content of the stalks from which they are growing. Genotypic and environmental factors influence sucker number. Several experiments were designed to identify environmental stimuli for suckering. Increased nitrogen, through application late in the growing season, was found to increase the numbers of suckers present. Manipulation of the light environment of main stalks showed that a change in light quality (but not necessarily quantity) stimulated suckering. Further work is underway to define how these environmental stimuli lead to sucker initiation and growth. Major scientific questions that should be addressed are: the identification of other potential environmental stimuli, how the signals are perceived and translated into sucker initiation, and why suckers have different morphology. The practical challenge is to combine the new information about sucker biology and develop from it strategies to alter agronomy and select new cultivars that results in decreased suckering.     

Rangewide molecular structuring in the Utah sucker (Catostomus ardens)

The Utah sucker (Catostomus ardens) is endemic to the Bonneville Basin and the upper Snake River drainage in western North America, and is thought to hybridize with the federally endangered June sucker (Chasmistes liorus mictus) in Utah Lake (Bonneville Basin). Here we describe the discovery of a major subdivision in Utah suckers (4.5% mitochondrial sequence divergence) between the ancient Snake River drainage and the Bonneville Basin. This boundary has not previously been recognized in Utah suckers based on morphologic variation, but has been recently described in two endemic cyprinids in the region. Populations in valleys east of the Wasatch Mountains in Utah clustered with the Snake River populations, suggesting that these valleys may have had an ancient hydrologic connection to the Snake River. We also found evidence of population isolation within the Bonneville Basin, corresponding to two Pleistocene sub-basins of the ancient Lake Bonneville. In contrast, we found no molecular evidence for deep divergence between Utah suckers and June suckers in Utah Lake or for a history of hybridization between divergent lineages in that population, although we recognize that demographic events may have obscured this signal. These findings suggest that the morphological differences between Utah and June suckers in Utah Lake may be the result of strong, and relatively recent, ecological selection. In summary, morphological and molecular characters seem to vary along different axes in different portions of the range of this taxon, providing an interesting system for studying the contributions of neutral and adaptive variation to species diversity.     

Segregation by species and size classes of rainbow trout,Salmo gairdneri,and Sacramento sucker,Catostomus occidentalis,in three California streams

Synopsis The hypothesis that Sacramento suckers, Catostomus occidentalis, compete with rainbow trout, Salmo gairdneri, for space in streams was examined by measuring microhabitat utilization of both species in three California streams. Two streams were similar in most respects except one contained only trout and one contained trout and a large population of suckers. The third stream, formed by the union of the first two, contained trout and a small population of suckers. The species overlapped in five of the six microhabitat variables measured: maximum depth, mean water column velocity, focal point velocity, surface water velocity, and substrate type. However, the species had strong vertical segregation; there was little overlap between species in focal point depth. Mean focal point velocities were also significantly different. Suckers roamed over and generally remained in contact with the bottom while trout held position in the water column. Microhabitat utilization by trout in the stream without suckers was similar to in the stream with a higher sucker density. Differences in microhabitat utilization by trout between the third stream and the other two was attributed to the larger size of the third stream. Both sucker and trout showed a similar within-species segregation of size classes - fish under 50 mm in length sought shallow water. Size-specific trends indicated ontogenic shifts in resource utilization which reduced overlap within species. These results suggest that competition for space between trout and suckers was not a major factor regulating microhabitat utilization of trout, although the possibility that larger suckers may displace small trout needs further study.     

日本棘隙吸虫体被超微结构的研究

本文报道扫描电镜观察下日本棘隙吸虫皮式、体棘和3种感觉乳突的形态和分布。     

Morphological and ultrastructural aspects of Branchiobdella pentodonta Whit. (Annelida,oligochaeta) suckers

The muscular system in the posterior sucker of Branchiobdella pentodonta Whit. has circular, longitudinal and radial fibers. In the anterior sucker, which has circular and longitudinal fibers, the muscle system is scarce. Concentric fibers are found around the mouth. In both suckers the glandular elements form voluminous complexes secreting mucus for attachment to the substrate. Suckers show neuromuscular junctions and three distinct types of neuroglandular junctions: one with typical neurosecretory granules, one with larger neurosecretory granules produced by cells located at the origin of the segmental nerves, and one with presynaptic vesicles. The second type is peculiar to the posterior sucker. A comparison is made between suckers of Branchiobdella and those of leeches.     

Variation in Scolex Morphology within and between Some Species of the Genus Proteocephalus Weinland (Cestoda, Proteocephala) with References to Strobilar Morphology

The scolex morphology of some Proteocephalus species from fish caught in fresh and brackish waters in Norway, as revealed by scanning and transmission electron microscopy and light microscopy, is described. The size and form of the fifth apical sucker proved to be a very pure character for identification. In spite of this, however, the scoleces of different species revealed specific characters which together with strobilar characters should be well suited for species discrimination and identification.     

头足类腕上吸盘和内壳扫描电镜观察

本文采用扫描电子显微镜对7种头足类腕上的吸盘及乌贼的内壳进行观察。结果表明,蛸科（Octopodidae）的短蛸（Octopus ocellatus）、真蛸（O. vulgaris）和长蛸（O. variabilis）腕的吸盘均无齿,乌贼科（Sepiidae）的日本无针乌贼（Sepia japonica）和金乌贼（S.esculenta）的腕和触腕吸盘上的角质环均有齿,且齿的形状不一样;虎斑乌贼（S. pharoanis）触腕的吸盘角质环光滑无齿,枪乌贼科(Loliginidae)的中国枪乌贼（Uroteuthis chinensis）腕上吸盘的角质环无齿。对３种乌贼的内壳的横切面进行扫描电镜观察,乌贼的内壳的背表面和腹表面的结构均是由文石和铺在上面的球型颗粒构成,但文石的形状和小球的排列存在种间差异,这些差异可以作为头足类的分类辅助特征。     

Revision of Gangesia (Cestoda: Proteocephalidea) in the Indomalayan Region: Morphology,Molecules and Surface Ultrastructure

Tapeworms of Gangesia Woodland, 1924 (Cestoda: Proteocephalidea) parasitic in freshwater fishes in the Indomalayan Region were critically reviewed. Evaluation of type specimens and newly collected materials from Bangladesh, Cambodia and India, as well as critical examination of extensive literature have shown that only the following four species, instead of 48 nominal species of Gangesia and Silurotaenia Nybelin, 1942 reported from this region (36 new synonymies proposed), are valid: Gangesia bengalensis (Southwell, 1913), type-species of the genus and most common parasite of Wallago attu (Siluridae), G. macrones Woodland, 1924 typical of Sperata seenghala (Bagridae), both species characterized by the possession of two circles of hooks on the rostellum-like organ and several rows of hooklets on the anterior margins of suckers; G. agraensis Verma, 1928 from W. attu (typical host), which has the scolex with only one circle of hooks and 1–3 incomplete rows of tiny hooklets on the suckers; and G. vachai (Gupta and Parmar, 1988) n. comb. from several catfishes, which possesses 4–6 circles of hooks and 5–11 rows of hooklets on the anterior half of suckers. Scolex morphology, including surface ultrastructure (microtriches), of all but one species (G. vachai) is described for the first time using scanning electron microscopy. A phylogenetic analysis based on the partial sequences encoding the large nuclear ribosomal subunit RNA gene has shown that three Indomalayan species, namely G. bengalensis, G. macrones and G. vachai, form a monophyletic group within Gangesia, whereas G. agraensis tends to form a clade with the Palaearctic species of the genus. A table with differential characters of all species from the Indomalayan Region is also provided together with a key to identification of genera of the subfamily Gangesiinae. The present study demonstrates that species of Silurotaenia do not occur in the Indomalayan region.     

胡杨（Populus euphratica）在额济纳绿洲三种生境内的根蘖繁殖特性

以额济纳荒漠绿洲胡杨根蘖幼苗为研究对象,通过对绿洲内胡杨分布的3种主要生境内根蘖幼苗的调查,得出如下结论。近年来绿洲内胡杨种群的更新几乎完全依赖于无性繁殖更新,种子萌发产生的幼苗在调查地内没有发现。河岸沙丘地及胡杨林下地的根蘖幼苗中,3龄幼苗所占比重最大,龄级越小的根蘖苗在群落中的数量越少;在河水漫灌后的林间空地,2龄幼苗的数量最多。水漫灌后的林间空地中根蘖幼苗密度显著大于其余两生境内根蘖幼苗密度,同时该生境内每段胡杨根系萌生出的不定芽也大于另两生境。随着龄级增加,未枯枝根蘖幼苗的比率逐渐减小。根蘖幼苗的萌发点分布于土层30cm以内的根系上,分布深度大于30cm的根系不能萌发出根蘖幼苗。     

Spatial organization of musculature in the Himasthla elongata cercaria (Trematoda: Echinostomatidae)

Somatic muscles (body-wall and "parenchyma" musculature), muscles of suckers, alimentary tract and excretory bladder of Himasthla elongata cercaria were investigated using fluorescent phalloidin labelling and confocal microscopy. The arrangement of body-wall muscles differs between the certain parts of cercarial body and appears to be the most complicated in the collar district. Among the body-wall musculature, we described U-shaped muscles, which have never been found previously in trematodes. Muscles of oral and ventral suckers are grouped into 6-7 independent layers. In some of those layers, they are arranged bilaterally, which contradicts the tradition to consider the sucker as radially symmetric.     

The function of the suckers of larval net-winged midges (Diptera: Blephariceridae)

1. Larval net-winged midges (Diptera: Blephariceridae) possess six ventral suckers that enable them to inhabit swift streams. Each sucker consists of a suction disc and a cavity with a piston. Large muscles are inserted within the piston, as well as at the base of the suction disc. This structure infers that both attachment and release of the sucker is achieved by vertical movements of the piston.
2. Live observations of blepharicerid larvae revealed that the sucker is indeed attached by an upward movement of the piston, but that the cavity is flooded when the sucker is released. The piston is lowered only at the end of a sucker 'step', expelling water from the cavity.
3. During foraging, the maxilla and the piston of the first sucker are moved synchronously, indicating that the first sucker functions as a holdfast thus facilitating grazing.
4. The adhesive forces, as well as the relative size of blepharicerid suckers, differ amongst species. They are highest in Hapalothrix lugubris and lowest in Liponeura cordata , which correlates with the hydraulic stress to which the larvae of these species are exposed in their preferred habitat. The balance between the efficiency of their retention structure and the hydraulic conditions of their preferred habitat defines a key dimension of their ecological niche.     

Effects of stand age and litter removal on the regeneration of Populus tremuloides

Abstract. The objective of this study was to demonstrate the regeneration by root suckering of Populus tremuloides (quaking aspen) along a successional sere at Lake Duparquet in NW Québec with six sites of different ages - 46, 74, 120, 143, 167 and 230 yr since last fire. In 1990, in each of the sites, one 20 m x 20 m plot was cut. In each plot, 4-m² plots were selected, both control and with litter removed. The number of suckers and their height were estimated for each plot in 1991, 1992 and 1993. Results showed that sucker production was not related to the number of P. tremuloides before the clearcut, but it was related to litter whereby increasing cover lowered the production. The number of suckers produced in the control plots did not show any significant difference along the successional sere for any of the three years. Scarified plots had more suckers than control plots, and this increase was more pronounced and significantly different for the younger stage. Regression analysis for the 1991 and 1992 data showed that sucker number in scarified plots decreased with increasing seral stage age. Mortality, after three growing seasons, was about 60% for all plots and was not affected by seral stage and treatment. We suggest that removal of litter in scarified plots exposed the roots to higher temperatures and caused wounds, both of which may have stimulated the production of suckers in young seral stages.     

Potentially handicapped but otherwise functional: Malformations in prey capture tools show no impacts on octopus life

Larval mortality is a keystone ecological factor for many benthic octopus since it mostly occurs before their settlement in the sea bottom as benthic juveniles. The literature had revealed that records of adult animals with morphological abnormalities (teratologies) are fewer in species with complex life cycle than in those with direct development. This is a direct consequence of the morphological, physiological, and development challenges that the transition from the larval to the adult morphology represents. During a routine fishing sample, we found an immature female horned octopus with additional buccal structures in two suckers of its ventral arms, likely rendering these suckers as inefficient. Based on the literature about the natural history of octopus, we provide evidence that these abnormalities were present at the moment of hatch. We evaluated the impact of the teratologies by comparing the shape of the buccal beaks and the trophic niche of the individual with five normal conspecifics. Although the beaks showed a different shape than normal individuals, the trophic niche was similar. Surprisingly, the teratological condition of the individual likely had no severe impacts on its life, even though it likely represents a handicap for its survival during its planktonic life. We also comment on other previous records from the literature of teratological adult octopus to highlight the amazing adaptive capacity of octopus to deal with challenging morphologies.     

Sensory innervation in the rim of the octopus sucker

Anatomical components of afferent innervation in the rim of the octopus sucker are described. In the sensory epithelium under the smooth cuticle two associated ciliated receptor cell-types (presumably chemosensitive) occur in clusters. A third ciliated receptor cell-type under the toothed cuticle may be a mechanoreceptor. A non-ciliated receptor cell-type of unknown function, under the toothed cuticle, is characterized by a microvillus-lined apical canal containing dense granular material. The axons of the latter two receptors go directly into large nerve tracts which nm through the infundibular muscle and on to the ganglion of the sucker. The axons of the first cell-types terminate on interneurons either in the base of the epithelium or below the epithelium. All the interneurons of the basal region of the epithelium migrate centripetally and develop into encapsulated interneurons. Within the epithelium, fine fibers provide collateral contact among cluster receptors. Collateral interaction among basal and encapsulated interneurons occur in the infundibular plexus. The microanatomy of the rim of the sucker suggests that chemosensory cues are funneled into the interneurons where they are concentrated into integrated signals, while other sensory input is probably sent directly to the ganglia of the sucker and/or arm.