Terminology and functional morphology of attachment structures in pelmatozoan echinoderms

Brett, Carlton E. 1981 12 IS: Terminology and functional morphology of attachment structures in pelmatozoan echinoderms. Lethaia . Vol. 14, pp. 343–370. Oslo. 1SSN 0024–1164.
Fixation on the seafloor is a fundamental, though little studied, aspect of the biology and distributional ecology of pelmatozoan echinoderms. Morphology of pelmatozoan attachment structures (holdfasts) and their component appendages is summarized herein and a detailed terminology proposed. Three major modes of fixation are differentiated in living and fossil stemmed echinoderms - primary, composite and secondary attachments - in addition to the direct thecal attachment in stemless forms. Specific holdfast morphologies, which characterize certain major taxa (families and orders) of pelmatozoans, may limit the available life-modes in these echinoderms. Significantly, numerous lineages of echinoderms have independently developed secondary attachment modes associated with the evolution of flexible tapered stem ends and/or true cirri. Cirri are rare in early Paleozoic pelmatozoans but appeared in most major groups by the late Paleozoic. * Pelmatozoa. attachment structures, holdfasts, terminology, functional morphology .     

Structure of the tarso-pretarsal chordotonal organ in the imago of Tineola bisselliella Humm. (Lepidoptera : Tineidae)

The tarso-pretarsal chordotonal organ in the imago of Tineola bisselliella (Lepidoptera .: Tineidae) includes 2 groups of scolopidia. (1) A basal group composed of 2 scolopidia, each with one neuron and one scolopidium with 3 neurons, whose dendrites present a typical structure of cilium; the dendrite becomes dense apically; it is a common characteristic for all the mechanoreceptive dendrites. (2) An apical scolopidium with 3 neurons, whose dendrites have the same size and are covered with a conical cap.The combination of single and triple dendrites is unusual in the limb chordotonal organs. The 2 groups of scolopidia are not in contact. The importance of the support structures and fixation structures is discussed. In the ciliary root region, peculiar desmosomes occur between the dendrite and the scolopale cell. For the apical scolopidium, one attachment cell is seen distally and it appears in close association with the articular membrane.     

The morphological and ontogenetic changes in the Protozoa during the transition to a sessile mode of life

The morphological adaptations of protozoans to sessile mode life and evolutionary changes in ontogeny are considered. There are main morphotypes of sessile protists: stalked organisms that attached to substrate by the extended base of body (basal disk), and unstalked organisms that are flatted on substrate. The origin of the morphotypes was independent in different taxa and involved nonhomologous structures. Adaptation to the sessile mode of life in the protists was connected with the progressive increase in the body size and intensity of organelle functions by polymerisation, subsequent division of function and change of functions. Evolution of adhesive organelles is characterised by growing intensity of their functions by allometric growth (usually without polymerisation), and in some cases with the subsequent division of functions and change of functions. The evolution manifests itself primarily in the organelles that provide interaction of cell with environment. The organelles that ensuring functioning of cell change due to correlations with the organelles of the first group. These two groups of organelles are similar to A.N. Sewertsoff's ecto- and endosomatic organs in multicellular organisms. The ontogeny of the sessile protists included three stages: formation of the migratory stage, distribution and choice of substrate and metamorphosis of the migratory stage after adhesion. As a rule there are no recapitulations on the first stage. The majority of structures tomotes or zoospores are inherited from the parent cell. Thus the present of some ancestral characteristics at the earlier stages of protistean ontogeny is display of the Baer's law. The main features of ontogeny evolution in sessile protists are the anaboly of the additional stages of life cycle, the displays of archallaxis or deviation during the migratory stage formation, and anaboly at the stage of buds morphogenesis after adhesion. At the last stage, the study of recapitulations is most perspective with the decision of phylogenetic problems in sessile protists.     

Attachment structures in Rhizotrochus (Scleractinia): macro‐ to microscopic traits and their evolutionary significance

Marine sessile benthic organisms living on hard substrates have evolved a variety of attachment strategies. Rhizotrochus (Scleractinia, Flabellidae) is a representative azooxanthellate solitary scleractinian coral with a wide geographical distribution and unique attachment structures; it firmly attaches to hard substrates using numerous tube‐like rootlets, which are extended from a corallum wall, whereas most sessile corals are attached by stereome‐reinforced structures at their corallite bases. Detailed morphological and constructional traits of the rootlets themselves, along with their evolutionary significance, have not yet been fully resolved. Growth and developmental processes of spines in Truncatoflabellum and rootlets in Rhizotrochus suggest that these structures are homologous, as they both develop from the growth edges of walls and are formed by transformation of wall structures and their skeletal microstructures possess similar characteristics, such as patterns of rapid accretion and thickening deposits. Taking molecular phylogeny and fossil records of flabellids into consideration, Rhizotrochus evolved from a common free‐living ancestor and invaded hard‐substrate habitats by exploiting rootlets of spines origin, which were adaptive for soft‐substrate environments.     

Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China

The Chengjiang Lagerstätte in the Lower Cambrian of South China yields a small, larva‐like arthropod, which was considered to be a protaspis of naraoiids by many authors. The discovery of a large number of well‐preserved specimens from many new localities has allowed the original study to be revised. The relatively large size, stable morphology and unusual structure of the appendages indicate that these specimens represent adults of a new arthropod, Primicaris larvaformis. The larva‐like outline is considered to have arisen by the heterochronic process of progenesis. In addition, this animal displays primitive aspects of bodyplan and limb morphology that suggest a basal position within arachnomorphs, or perhaps even arthropods, and the similarities to the Vendian arthropod‐like animal Parvancorina probably provide an evolutionary link between Vendian forms and Cambrian arthropods.     

Morphological changes and resource allocation of Zizania latifolia (Griseb.) Stapf in response to different submergence depth and duration

Plants around ponds, rivers and lakes are subjected to long-term partial or complete submergence. When they are flooded, water level affects the plants simultaneously with duration of submergence. Separate and interactive effects of water level and duration on the growth of the herbaceous perennial Zizania latifolia (Poaceae) were investigated by exposing the plants in greenhouse water tanks to submergence in different water depths and for different time-spans. The plants exhibited great shoot elongation upon submergence and prolonged flood duration, and the basal tiller number of the species decreased with higher water levels. Submergence treatment advanced the flowering date and increased the inflorescence number. Plant total biomass did not differ among all the treatments, while the root:shoot ratio decreased with increased water level, prolonged duration of submergence and their interaction. The high plasticity in morphology and shifts in reproductive strategy and biomass allocation enabled the Zizania plants to survive the compound effect of flooding height and duration. This may explain the occurrence of this species in habitats subjected to long-term flooding. The results obtained in this experiment will contribute to understanding the impact of flooding dynamics on plants and the ways of adaptation responses to prolonged waterlogging.     

Fedomia mikhaili—A new spicule-bearing organism of sponge grade from the Vendian (Ediacaran) of the White Sea, Russia

The Vendian fossil locality on the Solza River is one of the most productive in the White Sea Region. The fossiliferous deposits belong to the Upper Vendian Verkhovka Formation correlated lithologically with strata dated at 558 ± 1 to 555.3 ± 0.3 Ma in adjacent regions. This locality is characterized by yielding more than 10 well-represented Ediacaran genera. Elongated imprints of Fedomia mikhaili, n. gen. et n. sp., are common at the Solza River fossil locality. The new taxon is characterized by its six- to eight-rayed star-shaped concave structures with diameters of 2–5 mm. The organism can be reconstructed with sacciform thin-walled body attached basally to the substrate. The star-shaped structures may be spicules that appear to be rather flexible than rigid. The new genus, Fedomia, shows some similarities to Eiffelia Walcott, 1920 from the Middle Cambrian of British Columbia, and thus is interpreted as an organism of sponge grade. The new taxon could represent an additional phylogenetic link between the Ediacaran and Cambrian worlds.     

Crystal structures of the key anaerobic enzyme pyruvate:ferredoxin oxidoreductase, free and in complex with pyruvate

Oxidative decarboxylation of pyruvate to form acetyl-coenzyme A, a crucial step in many metabolic pathways, is carried out in most aerobic organisms by the multienzyme complex pyruvate dehydrogenase. In most anaerobes, the same reaction is usually catalyzed by a single enzyme, pyruvate:ferredoxin oxidoreductase (PFOR). Thus, PFOR is a potential target for drug design against certain anaerobic pathogens. Here, we report the crystal structures of the homodimeric Desulfovibrio africanus PFOR (data to 2.3 A resolution), and of its complex with pyruvate (3.0 A resolution). The structures show that each subunit consists of seven domains, one of which affords protection against oxygen. The thiamin pyrophosphate (TPP) cofactor and the three [4Fe-4S] clusters are suitably arranged to provide a plausible electron transfer pathway. In addition, the PFOR-pyruvate complex structure shows the noncovalent fixation of the substrate before the catalytic reaction.     

Microbial stabilization of sediments in a recent Salina, Lake Aghormi, Siwa Oasis, Egypt

Stabilization of sediments by microbial mats and biofilms were studied in detail in Lake Aghormi, Siwa Oasis, Egypt. The study has shown that microbial mat assemblages, particularly filamentous cyanobacteria, with their extracellular polymeric substances (EPS) are capable of effectively stabilizing sediments. The microbial mats in the siliciclastic environments of Lake Aghormi display distinctive sedimentary structures (microbially induced sedimentary structures), including multidirected ripple marks, microbial patches, petee structures, erosional remnants and pockets, and gas domes. Scanning electron microscopy study of the sediment surface colonized by cyanobacteria revealed that filamentous types are the most effective stabilizing organisms. Filamentous cyanobacteria and their EPS construct a network, interweave depositional grains of the sediment surface, envelope the particles, and glue them together. The studied biofilm is so thick forming a spider-web structure that totally coat the particles in such a way the morphology of the particles is masked.     

Antennal sensory organs of a troglobitic coleoptera,Speonomus hydrophilus jeannel (Catopidae): An ultrastructural study by chemical fixation and cryofixation

The ultrastructure of 12 antennal sensory organs of the cavernicolous beetle Speonomus hydrophilus (Coleoptera : Catopidae) after chemical fixation or cryofixation is described. This work helps us to compare the results of the 2 types of fixation; cryofixation is more difficult and hazardous, but when it succeeds, the cell preservation is better than with conventional chemical fixation; the cell shapes are more regular and rounded, and some details, such as microtubule subunits, can be visible only after cryofixation. On the basis of these ultrastructural features, the sensory organs are functionally classified. The structures of these organs are compared in epigean and troglobitic Coleoptera.     

Tarsal attachment devices of the southern green stink bug Nezara viridula (Heteroptera: Pentatomidae)

Based on analyses with cryo‐scanning and transmission electron microscopy, the present study reports on the morphology and ultrastructure of the attachment structures of the green stinkbug Nezara viridula L. (Heteroptera: Pentatomidae), a cosmopolitan pest of different crops in most areas of the world. In addition, the presence and distribution of large proportions of the elastic protein resilin in these structures was revealed by confocal laser scanning microscopy. The attachment structures of each leg comprise two sclerotised claws, a pair of smooth flexible pulvilli and a hairy adhesive pad located at the ventral side of the basitarsus. No sexual dimorphism is evident. Contact areas of resting individuals on a smooth surface show that N. viridula creates contact to the substrate with the ventral surface of (a) the distal portions of the pulvilli, (b) the setae of the hairy adhesive pad, (c) the two paraempodia representing mechanosensory setae, and (d) the tips of the claws. Each pulvillus is a sac‐like structure formed by complex cuticular layers that vary in their structure and resilin content. The dorsal side consists of sclerotised chitinous material, while the ventral cuticle consists mainly of resilin and shows a very thin epicuticle and a thick exocuticle. The setae of the hairy adhesive pad are pointed and socketed. They exhibit a pronounced longitudinal gradient in the material composition, with large proportions of resilin being present in the setal tips. In most of these setae, especially in those of the distal‐most part of the pad, also a transverse gradient in the material composition is visible.     

Bioaccumulation in Miscanthus sacchariflorus grown on cadmium-contaminated sediments: a comparative study between submerged and non-submerged environments

Bioaccumulation of heavy metals in aquatic plants is significantly affected by hydrological regime and therefore the accumulation and translocation of cadmium in five organs—panicle, leaf, stem, root, and bud—of an emergent plant (Miscanthus sacchariflorus) were compared between the submerged environment and non-submerged environment. In the submerged condition, the cadmium concentration was higher in the panicle and leaf than in the stem, root, and bud. Cadmium concentration in the root exhibited a positive regression with cadmium concentration in the sediment. However, cadmium concentration in the panicle, leaf, stem, and bud exhibited no significant regression with cadmium concentration in the sediment. In the non-submerged environment, the cadmium concentration was higher in the below-ground organs than in the aboveground organs. The mean bioaccumulation coefficient in the 24 investigated plots in the submerged environment was higher than that in the 20 and 40?mg kg^?1 cadmium treatments in the non-submerged environment. The mean translocation factor in the submerged environment was nine times higher than that in non-submerged environment. These results indicate that submergence enhanced cadmium bioaccumulation in the aboveground organs and that this plant can be used to remove heavy metals from polluted rivers and lakes.     

A Novel Basal Body Protein That Is a Polo-like Kinase Substrate Is Required for Basal Body Segregation and Flagellum Adhesion in Trypanosoma brucei

The Polo-like kinase (PLK) in Trypanosoma brucei plays multiple roles in basal body segregation, flagellum attachment, and cytokinesis. However, the mechanistic role of TbPLK remains elusive, mainly because most of its substrates are not known. Here, we report a new substrate of TbPLK, SPBB1, and its essential roles in T. brucei. SPBB1 was identified through yeast two-hybrid screening with the kinase-dead TbPLK as the bait. It interacts with TbPLK in vitro and in vivo, and is phosphorylated by TbPLK in vitro. SPBB1 localizes to both the mature basal body and the probasal body throughout the cell cycle, and co-localizes with TbPLK at the basal body during early cell cycle stages. RNAi against SPBB1 in procyclic trypanosomes inhibited basal body segregation, disrupted the new flagellum attachment zone filament, detached the new flagellum, and caused defective cytokinesis. Moreover, RNAi of SPBB1 confined TbPLK at the basal body and the bilobe structure, resulting in constitutive phosphorylation of TbCentrin2 at the bilobe. Altogether, these results identified a basal body protein as a TbPLK substrate and its essential role in promoting basal body segregation and flagellum attachment zone filament assembly for flagellum adhesion and cytokinesis initiation.     

Development of algal-foraminiferal-coral reefs in the Lower Carboniferous of Furness, northwest England

Most carbonate buildups of Dinantian age are mud-mounds lacking direct evidence of abundant framework organisms. This contribution describes apparently unique structures containing abundant framebuilding organisms interpreted as true reefs. They occur in the Red Hill Oolite, part of the Carboniferous Limestone succession in the Furness area of northwest England. Reefs were initiated by the attachment of numerous Syringopora colonies to a firm substrate. Encrusting organisms, dominantly the supposed foraminifcr Aphralysia, colonised sediment and corallite surfaces leading to the development of a rigid framework. Thrombolites also assisted in the establishment of bindstone textures. During the later stages of reef growth, Syringopora became less common and its place in the reef was taken by upright, branching growths of solenoporoid algae. Rapid sedimentation and subsidence resulted in reefs with near vertical sides, but little topographic expression on the sea-floor during growth. The occurrence of these reefs cannot be attributed to any single environmental factor but probably resulted from an unusual combination of favourable circumstances. D Calcareous algae, Carboniferous, corals, Dinantian, foraminifera, reefs, thrombolites.     

Complex tibial organs in fore-, mid-, and hindlegs of the bushcricket Gampsocleis gratiosa (Tettigoniidae): Comparison of morphology of the organs

The structure of the complex tibial organs in the fore-, mid-, and hindlegs of the East Asian bushcricket Gampsocleis gratiosa (Tettigoniidae, Decticinae) is described comparatively. In each leg the tibial organs consist of three scolopale organs: the subgenual organ, the intermediate organ, and the crista acoustica. Only in the forelegs are the tibial organs differentiated as tympanal organs, and sound transmitting structures (acoustic trachea, tympana, and tympanal covers) are present. The morphology of the tracheae in the mid- and hindlegs is significantly different from that found in the forelegs. The number of scolopidia in the subgenual organ is highest in the midleg and lowest in the foreleg; in the intermediate organ the number is also highest in the midleg, and the fore- and hindleg contain 40% fewer scolopidia. In the crista acoustica, the number of scolopidia decreases from, the fore- to the mid- and hindlegs. The morphology and the dimensions of the scolopidia and the attachment structures within the crista acoustica of the mid- and hindlegs differ strongly from those in the foreleg. The results indicate that, in addition to the presence of a sound transmitting system, the specific differentiations within the crista acoustica are important for the high auditory sensitivity of the tibial organs in the forelegs. © 1994 Wiley-Liss, Inc.     

三峡水库长期水淹条件下耐淹植物甜根子草的资源分配特征

为研究历经三峡水库长期水淹的驯化后,不同海拔高程之间相同种源的甜根子草的生物量分配特征是否发生改变,进而探讨该物种对水淹胁迫表现出的适应性进化特征,试验于2008年初选育相同种源的甜根子草同龄幼苗栽植于三峡水库消落区甜根子草种植试验示范区,并考察了2012年、2013年不同海拔高程甜根子草植株的形态和生物量特征。试验共设置3个海拔高程,即水淹高程168、172m和不受水淹对照高程176m。试验结果表明:(1)较低高程的甜根子草植株较矮小细弱,168m高程的甜根子草植株主茎长和主茎基径显著低于对照176m高程(P0.05);平均节间长度随高程的降低而缩短;与之相反,主茎长/主茎基径随高程的降低而增大。(2)甜根子草的叶片厚度、叶片长/叶片宽、叶片长/叶鞘长均随海拔高程的降低而减小;与之相反,比叶面积随高程的降低而增大。(3)水淹前,甜根子草近端成熟节间的质量密度随高程的降低而增大;水淹后,其地上存活茎段基部成熟节间的质量密度在各高程之间无显著差异(P0.05)。以上研究结果表明,甜根子草历经三峡水库长期水淹的驯化后,生物量分配特征在不同海拔高程之间发生了改变,表现出了相应的驯化特征。相较于高高程的甜根子草植株而言,低高程的植株生长缓慢,采取低株高下的高向生物量投资策略;对叶的物质投资大部分分配到叶面积的增加、叶鞘的伸长生长和叶片的直立生长上,以加强植株的光合生产。     

Asymmetrical soft-sediment deformation structures triggered by rapid sedimentation in turbiditic deposits (Late Miocene, Guadix Basin, southern Spain)

Summary Soft-sediment deformation structures in Tortonian turbiditic deposits of the Guadix Basin (southern Spain) have been described. The most common structures are asymmetrical pillow structures and elongated sets of loadcasts. The structures are metric in scale and have been interpreted as the result of liquefaction and/or fluidization processes triggered by the rapid sedimentation of single high concentration turbidites. Final morphology of soft-sediment deformation structures is related to two main driving force systems: unstable density gradient and lateral shear stress. The latter is probably induced by the downslope component of the sediment weight. The asymmetry of deformational structures (in horizontal and vertical cross-section) allows a clarification of the relationship between morphology of deformation and direction of lateral shear stress: this relationship seems ambiguous and confused in the literature. The interpretations both of deformation mechanism and trigger agent have been supported with:-field analyses;-calculations on the liquefaction processes induced by rapid sedimentation;-qualitative models in laboratory.     

Insect walking techniques on thin stems

The attachment ability of insects on surfaces are associated not only with the micro- and nanostructure of the adhering part of an attachment device, but also with the global scale kinematics responsible for contact formation and release. In the present study, the locomotory techniques of several representatives of insects from four different orders (Orthoptera, Heteroptera, Coleoptera, and Hymenoptera), possessing different types of attachment structures, are described. The study is based on video recordings of insects walking on a flat surface and on cylindrical rods of various thickness, imitating plant stems. Attachment devices of tarsi and pretarsi were visualized using Scanning Electron Microscopy. The results show a different manner in the use of adhesive structures on substrates with various curvatures. Insects bearing attachment pads on proximal tarsomeres usually touch flat and curved substrates using all tarsomeres, whereas insects with their attachment devices on the distal tarsomeres usually walk on flat surfaces using the distal tarsomeres of the overextended tarsus. On substrates, with diameters comparable to or larger than the tarsus length, insects walk above the stem by clasping the stem with the bent tarsi. On thin stems, insects clasp the stem between their tarsi and hang under the stem. Thus, on thin and thick rods, forces applied to attachment organs act in opposite directions. There are two methods of leg positioning for walking on a rough flat substrate. In the first case, the tarsus is straightened and the rough substrate is gripped between the claws and the proximal complex of attachment devices (tarsal euplantulae, fossulae spongiosa, and terminal spurs of tibiae). In the second case the tibia does not touch the substrate; the insect is supported only by distal tarsomeres. The tarsus is in an overextended condition. On rods, with diameters comparable to or larger than the tarsus length, insects walk by clasping the stem with the bent tarsi. This posture is characteristic for the majority of insects independent of the tarsal position they normally use while walking on a plane. If the rod’s diameter is smaller than the tarsus length, walking insects usually clutch it between contralateral tarsi. Using such a posture they are supported by interlocking or by strong friction, generated by attachment devices of the proximal tarsomeres, and do not use attachment devices of the pretarsus. Contact with the substrate is reinforced due to the coordinated contralateral clutch using all supporting legs. It is concluded that the use of different types of attachment structures correlates with locomotory techniques. Handling Editor: Heikki Hokkanen     

Evolution of sensory structures in basal metazoa

Cnidaria have traditionally been viewed as the most basal animalswith complex, organ-like multicellular structures dedicatedto sensory perception. However, sponges also have a surprisingrange of the genes required for sensory and neural functionsin Bilateria. Here, we: (1) discuss "sense organ" regulatorygenes, including; sine oculis, Brain 3, and eyes absent, thatare expressed in cnidarian sense organs; (2) assess the sensoryfeatures of the planula, polyp, and medusa life-history stagesof Cnidaria; and (3) discuss physiological and molecular datathat suggest sensory and "neural" processes in sponges. We thendevelop arguments explaining the shared aspects of developmentalregulation across sense organs and between sense organs andother structures. We focus on explanations involving divergentevolution from a common ancestral condition. In Bilateria, distinctsense-organ types share components of developmental-gene regulation.These regulators are also present in basal metazoans, suggestingevolution of multiple bilaterian organs from fewer antecedentsensory structures in a metazoan ancestor. More broadly, wehypothesize that developmental genetic similarities betweensense organs and appendages may reflect descent from closelyassociated structures, or a composite organ, in the common ancestorof Cnidaria and Bilateria, and we argue that such similaritiesbetween bilaterian sense organs and kidneys may derive froma multifunctional aggregations of choanocyte-like cells in ametazoan ancestor. We hope these speculative arguments presentedhere will stimulate further discussion of these and relatedquestions.