Valve Margin Research Articles

Common regulatory networks in leaf and fruit patterning revealed by mutations in theArabidopsis ASYMMETRIC LEAVES1gene

Carpels and leaves are evolutionarily related organs, as the former are thought to be modified leaves. Therefore, developmental pathways that play crucial roles in patterning both organs are presumably conserved. In leaf primordia of Arabidopsis thaliana, the ASYMMETRIC LEAVES1 (AS1) gene interacts with AS2 to repress the class I KNOTTED1-like homeobox (KNOX) genes BREVIPEDICELLUS (BP), KNAT2 and KNAT6, restricting the expression of these genes to the meristem. In this report, we describe how AS1, presumably in collaboration with AS2, patterns the Arabidopsis gynoecium by repressing BP, which is expressed in the replum and valve margin, interacts in the replum with REPLUMLESS (RPL), an essential gene for replum development, and positively regulates the expression of this gene. Misexpression of BP in the gynoecium causes an increase in replum size, while the valve width is slightly reduced, and enhances the effect of mutations in FRUITFULL (FUL), a gene with an important function in valve development. Altogether, these findings strongly suggest that BP plays a crucial role in replum development. We propose a model for pattern formation along the mediolateral axis of the ovary, whereby three domains (replum, valve margin and valve) are specified by the opposing gradients of two antagonistic factors, valve factors and replum factors, the class I KNOX genes working as the latter.

Phylogenetic relationships of coexisting Heterocypris (Crustacea, Ostracoda) lineages with different reproductive modes from Lampedusa Island (Italy)

Coexisting bisexual and unisexual populations of individuals belonging to the genus Heterocypris are found in ephemeral freshwater ponds on the island of Lampedusa (Pelagie Islands, Italy). Different reproductive modes were associated with a peculiar morphological trait: a lamella hyalina on the posterior margin of the left valve was observed in amphimictic females, a feature missing in apomictic females. In order to clarify the phylogenetic relationships among taxa with different morphological traits and reproductive modes, we used four polymorphic enzyme loci (GPI, ICD2, MPI and PGM) and mitochondrial DNA 16S ribosomal sequences. We identified three main evolutionary units that showed a combination of morphological and reproductive characteristics: (1) amphimictic females of H. barbara with a lamella hyalina, according to the typical feature of the species, and apomictic females of H. barbara without lamella that are sympatric in one temporary pond; (2) apomictic females of H. incongruens without a lamella, as typical of the species; (3) apomictic females without a lamella, living in sympatry with H. barbara, but characterised by a high genetic diversity from both H. incongruens and H. barbara. We discuss the possible origin of apomictic lineages as a result of independent transition episodes to apomixis from different sexual ancestors. Time of divergence reflected the genetic differentiation within and among multiple ancestors and different possible routes to parthenogenesis.

Contribution of external morphology in solving a species complex: The case of Prorocentrum micans, Prorocentrum gracile and Prorocentrum sigmoides (Dinoflagellata) from the Mexican Pacific Coast

SUMMARY Prorocentrum micans Ehrenberg, Prorocentrum gracile Schütt and Prorocentrum sigmoides Böhm are planktonic dinoflagellates present along the Mexican Pacific coast. These three species can be treated as a species complex because of similar shape and size. The taxonomy of the group has been based on cell form and size, the presence or absence of apical spines and the ornamentation of the main valves. A morphometric study of samples from 11 locations from the Mexican Pacific coast and from laboratory cultures is presented. The presence of a mucron (small tooth) in the antapical region of P. gracile and P. sigmoides morphotypes was recorded for the first time. Morphotypes were defined using as the main criteria the size of pores in relation to the size of the depressions on the cell’s surface, the presence of a mucron in the posterior end of the cell and the shape of the valve margins. Results show a clear distinction between the morphotype P. micans and the P. gracile–P. sigmoides. P. sigmoides is suggested to be a junior synonym of P. gracile as no convincing characters could distinguish these two morphotypes.

LEMNICOLA HUNGARICA (GRUNOW) ROUND & BASSON FROM SOUTHERN BRAZIL: ULTRA-STRUCTURE, PLASTID MORPHOLOGY AND ECOLOGY

Lemnicola hungarica (Grunow) Round & Basson taken from brackish and freshwater environments in Southern Brazil was studied for its ultra-structure details, plastid morphology, and ecology. Previously undescribed details are given: a ring of elongated areolae on the mantle of L. hungarica and its plate-like plastid that lies on the araphid valve where it reaches the valve margin and curves itself towards the raphid valve. Lemnicola hungarica thrives in truly freshwater conditions where conductivity values are below 300 μS/cm, and where floating plants are also present.

Sex in a cyclical parthenogen: mating behaviour of Chydorus sphaericus (Crustacea; Branchiopoda; Anomopoda)

Summary1. We describe the interactions during mating in Chydorus sphaericus, a cyclical parthenogenetic anomopod. Mating behaviour is more complex than previously assumed, with evidence for a diffusible chemical to which males react at the onset of mating, for reproductive isolation, and for postcopulatory mate guarding.2. During mating, the male and female form a ‘mating cross’ that may be maintained for several hours, while copulation itself typically lasts less than a minute. Furthermore, males invariably attach to the right valve of females. Copulation involves intromission of the postabdomen between the valves, so that the gonopores approach the left ovarium.3. This behaviour is reflected in the morphology of both sexes: males have a specialised anterior valve margin, postabdomen, first limb and rostrum, under selective pressure for successful mate guarding and copulation, while gamogenetic females have asymmetric ovaries, and a species‐specific setulation of the valves. Males of the structurally related Chydorus ovalis react to the presence of C. sphaericus, but fail to dock to females, suggesting a lock‐antilock element in the reproductive isolation of both species.4. The morphological and ethological adaptations in C. sphaericus suggest that there is a strong selective pressure on mating behaviour in this cyclical parthenogen and specifically towards the formation of the ‘mating cross’.

Structure and formation of the adventitious tube of the Japanese watering‐pot shell Stirpulina ramosa (Bivalvia, Anomalodesmata, Clavagellidae) and a comparison with that of the Penicillidae

Abstract. Stirpulina ramosa is the only extant endobenthic representative of the Clavagellidae and is restricted to the waters of Japan. A single intact adventitious tube of this species has been obtained and its structure is described. The right valve is 16 mm long and located within the adventitious tube. It has an opisthodetic ligament located on resilifers. There are anterior and posterior adductor muscle scars, a thick pallial line, and pallial and pedal gape (right valve only) sinuses. The left shell valve is but 9 mm long and is united into the fabric of the adventitious tube via the intermediary of a shelly saddle. Internally, only the anterior adductor muscle scar and a small element of the pallial line scar are identifiable on the left valve. The posterior adductor and the rest of the pallial line scar (including a pallial sinus) are, remarkably, located on the adventitious tube beyond the shell valve margin. The adventitious tube of S. ramosa is formed in a manner wholly dissimilar from that of Brechites vaginiferus (Penicillidae). In B. vaginiferus, the tube is secreted as a single entity from the general outer mantle surface, including the siphons, covering the body. As a consequence, both shell valves are incorporated into the structure of the tube and the watering pot is bilaterally symmetrical. In S. ramosa, the tube and watering pot are secreted from the mantle margin and surface surrounding and extending from the left shell valve, so that only the left valve is incorporated into its structure. A dorsally derived mantle element is progressively extended over to the right side of the body, meeting a ventrally derived counterpart that passes beneath it, forming a pleat in the calcareous structure of the right side of the tube that they secrete. This pleat extends into the complex of watering‐pot tubules and forms the pedal gape. The watering pot is thus Ω shaped. The ventrally derived mantle element forms a sinusoidal crest on the right‐hand base of the watering pot, creating a pedal gape sinus scar on the right valve. The Clavagellidae radiated widely in the Mesozoic, leaving behind a rich fossil record for Stirpulina. Only S. ramosa, however, has survived until the present. In contrast, the Cenozoic Penicillidae has a poor fossil record, but there is a rich variety of extant endobenthic watering‐pot shells. It has been argued hitherto that the two families represent a remarkable example of convergent evolution. In view of the success of the Penicillidae and thus the endobenthic, tube‐dwelling lifestyle, however, it is hard to understand why Stirpulina has largely died out—even S. ramosa being known by but one or two specimens. A study of the anatomy of S. ramosa might one day answer this question.

The INDEHISCENT protein regulates unequal cell divisions in Arabidopsis fruit

INDEHISCENT (IND) is one of the major factors of a transcriptional network regulating fruit dehiscence in Arabidopsis thaliana. However, partially owing to the lack of knowledge on the morphogenic process at the junction between the valve margin and the replum, the cellular processes regulated by IND have not been elucidated. Here we report the identification of a zone of seven layers of cells relevant to silique dehiscence, and the isolation and characterization of a transposon-enhancer-trap-generated null allele of IND, ind-6. Light microscopy analysis revealed that the beta-glucuronidase from the enhancer trap was expressed in a seven-layer zone, encompassing the valve margin and the replum border. Electron and light microscopy analyses also revealed that unequal cell divisions actively occurred at stage 15 in the seven-layer zone in wild type (WT), which apparently produced the distinct cell types in the WT seven-layer zone. In contrast, siliques of ind-6 at stage 15 did not exhibit such cell divisions but fewer cells and defective cytokinesis, hence lacking the distinct cell types in the seven-layer zone. These results demonstrate a cellular mechanism for establishing the distinct cell types and a role for IND in regulating the unequal cell divisions in the seven-layer zone.

(98) Molecular Analysis of Abscission Layer Activation in Apple Fruit Pedicels

Abscission of leaves, floral organs, and fruit is a developmentally and environmentally regulated process initiated in specialized thin layers of cells within abscission zones (AZs). Very little is known about early molecular events that drive abscission, especially of fruit. Commercial apple production relies on the use of flower and fruit abscission-promoting and -inhibiting compounds to enhance fruit quality, control preharvest fruit drop, and maintain consistent annual bearing. The success of chemical treatments is strongly influenced by numerous factors, including environment, genotype, developmental stage of the fruit, and physiological state of the tree. Toward developing improved strategies for regulating fruit abscission, we carried out transcriptional profiling of competent-quiescent and activated abscission layers. We found that a decisive event in the sequential process of abscission layer development is the transcriptional activation of the MdPEL1 gene, encoding a plant pectate lyase protein and potentially involved in the degradation of the middle lamella of adjacent abscission layer cells. Additionally, regulatory elements of at least 12 homologous pectate lyase genes in Arabidopsis thaliana were found to direct expression in floral AZs and in dehiscence zones along valve margins, suggesting that these genes have evolutionary conserved function. This work identifies a novel role for pectate lyases in plants. Furthermore, many abscission-related genes identified in this study are being used to track biochemical and regulatory pathways that participate in abscission in response to chemical treatments or environmental effects.

STUDIES ON THE DIATOM UROSOLENIA ROUND & CRAWFORD (RHIZOSOLENIOPHYCIDEAE) PART 1. NEW AND RE-CLASSIFIED SPECIES FROM SUBTROPICAL AND TROPICAL FRESHWATERS

Populations of the delicate planktonie freshwater centric diatom genus Urosolenia from several lentic freshwater environments in Africa, Asia, Australia and South America were studied with both LM and SEM. The classical morphological features, e.g. frustule size, shape of the valve, length and positioning of the process and number of girdle bands per unit length, were re-evaluated and additional previously overlooked valve structures were identified as useful for species differentiation. Although fine resolution LM revealed the most important elements of the spine-like process, the SEM showed additional ultrastructural features on the frustule surface and terminal ends as well as structures on the valve margins not resolved in detail by LM. A few new morphological terms are proposed mainly for terminal ends of the process (ligula, seta), the process shaft (poroids, protrusions, ribs) and the valve margin. Using these LM and SEM observations, the genus description is emended and two species previously described earlier under Rhizosolenia are transferred into the recently established genus Urosolenia (U. victoriae (Schroeder) Kling & Rott comb. nov. from Lake Victoria Africa and U. brauni (Hustedt) Rott & Kling comb. nov. from Jurucui Brasil). Three new Urosolenia species are described, two from Australia (U. brevispinosa Kling, McGregor & Rott sp. nov., U. parva Kling, Rott & McGregor sp. nov.), and one from Sri Lanka (U. diademata Rott & Kling sp. nov.). Type localities including ecological data of the environments are listed and a key for identification of the named species is given.

Effect of TBT on Ruditapes decussatus juveniles

The effects of sublethal concentrations of tributyltin (TBT) on growth of juvenile clams Ruditapes decussatus were determined during exposure to TBT concentrations of 50, 100 and 250 ng l −1 (as Sn) for a period up to two years. Length and weight of clams increased continuously in all treatments throughout the experimental period, and, overall, rates were not significantly influenced by TBT exposure, although final length and weight were inversely related to increasing TBT concentration. Juvenile R. decussatus therefore appear to be less sensitive to TBT than larval stages. Some juveniles exposed to TBT developed abnormal shell growth, laterally, changing the typical flattened shape of clams into a more “rounded” form. This characteristic was more visible in the anterior margins of valves than posteriorly, and mainly observed in clams exposed to TBT at 50 ng l −1 (as Sn).

A genetic framework for fruit patterning inArabidopsis thaliana

In the model plant Arabidopsis thaliana, the establishment of organ polarity leads to the expression of FILAMENTOUS FLOWER (FIL) and YABBY3 (YAB3) on one side of an organ. One important question that has remained unanswered is how does this positional information lead to the correct spatial activation of genes controlling tissue identity? We provide the first functional link between polarity establishment and the regulation of tissue identity by showing that FIL and YAB3 control the non-overlapping expression patterns of FRUITFULL (FUL) and SHATTERPROOF (SHP), genes necessary to form stripes of valve margin tissue that allow the fruit to shatter along two defined borders and disperse the seeds. FIL and YAB3 activate FUL and SHP redundantly with JAGGED (JAG), a gene that also promotes growth in organs, indicating that several pathways converge to regulate these genes. These activities are negatively regulated by REPLUMLESS (RPL), which divides FIL/JAG activity, creating two distinct stripes of valve margin.

Biology and functional morphology of a new species of endolithic Bryopa (Bivalvia: Anomalodesmata: Clavagelloidea) from Japan and a comparison with fossil species of Stirpulina and other Clavagellidae

Abstract. A new species of Clavagellidae, Bryopa aligamenta, from Okinawa, Japan, is described. The species is endolithic in living corals, with the left valve cemented to the crypt wall, as in all clavagellids. The free right valve exhibits an unusual growth pattern, with commarginal lines seemingly arising from the posterior valve margin and extending towards the anterior. This results from: (i) progressive anterior erosion of the umbones, probably as a consequence of the boring process; (ii) the apparent migration posteriorly, as the umbones are eroded, of the dorso‐ventral growth axis of the shell; and (iii) enhanced posterior inter‐commarginal growth. Unlike other clavagellid genera and species, however, there is no discernible primary ligament, at least in the adult. It is possible, however, that if a juvenile ligament were present (as in B. lata), it too would be lost as a consequence of antero‐dorsal erosion during boring. To retain valve alignment in the absence of a primary ligament, and possibly upon reaching an adult size, the mantle lays down alternating layers of calcium carbonate and proteinaceous periostracum onto the interior surface of the shell to thicken it, most noticeably marginally and, especially, posteriorly. The two valves are united dorsally, therefore, by thin layers of periostracum that probably exert a minimal opening force. B. aligamenta is, however, further characterised by large adductor, pallial, and siphonal retractor muscles so that the entire animal is encased tightly within an internally strengthened shell within a crypt. Movement must be minimal, blood being pumped into pallial haemocoels to push open the valves and extend the siphons. Despite a suggestion to the contrary, Bryopa is retained in the Clavagellidae, its unusual growth processes resulting from an endolithic life style within living corals.The fossil clavagellid Stirpulina bacillus, from the Pliocene/Pleistocene of Palermo, Sicily, Italy, was, unlike Bryopa aligamenta and other clavagellids, endobenthic, with a long adventitious tube and anterior watering pot superficially similar to species of Penicillidae, another family of the Clavagelloidea. Furthermore, as in all clavagellids only the left valve is fused into the fabric of the tube, the right being free within it. In all penicillids, both valves are fused into the fabric of their tubes. The watering pots of the fossil S. coronata, S. vicentina, and S. bacillus, moreover, are formed in a different manner to that of penicillids, by progressive encasement of the right valve inside the tube. In penicillids, the tube is secreted in a single event from the general mantle surface and the incorporation of both valves into its fabric. The constituent genera of the Clavagellidae thus constitute an example of parallel evolution with members of the Penicillidae.

Pod shatter‐resistant Brassica fruit produced by ectopic expression of the FRUITFULL gene

Arabidopsis has proven to be extremely useful as a reference organism for studies in plant biology, and huge efforts have been employed to unravel various mechanisms of Arabidopsis growth. A major challenge now is to demonstrate that this wealth of knowledge can be used for global agricultural and environmental improvement. Brassica species are closely related to Arabidopsis and represent ideal candidates for model-to-crop approaches as they include important crop plants, such as canola. Brassica plants normally disperse their seeds by a pod-shattering mechanism. Although this mechanism is an advantage in nature, unsynchronized pod shatter constitutes one of the biggest problems for canola farmers. Here, we show that ectopic expression of the Arabidopsis FRUITFULL gene in Brassica juncea is sufficient to produce pod shatter-resistant Brassica fruit and that the genetic pathway leading to valve margin specification is conserved between Arabidopsis and Brassica. These studies demonstrate a genetic strategy for the control of seed dispersal that should be generally applicable to diverse Brassica crop species to reduce seed loss.

High frequency and severity of breakage-induced shell repair in western Atlantic Pinnidae (Bivalvia)

High frequency and severity of breakage-induced shell repair in western Atlantic Pinnidae (Bivalvia)

Two new species ofDiaphanosomaFischer, 1850(Crustacea: Branchiopoda: Cladocera) from the United States

Two new species of the genus Diaphanosoma, D. oligosetum and D. dorotheae, from Louisiana and North Carolina respectively, are described. The former species has large head with protruding dorsal part, large lanceolate spine on the basipodite's distal outer end, an extremely reduced number of antennal setae, up to six in adult specimens, and unique armament of valve margin. On the whole, it shows the pronounced combination of primitive and specialized morphological traits. D. dorotheae is a member of D. brachyurum species group differing from its other known representatives in presence of a small but very conspicuous spine on the end of proximal segment of antennal exopodite and a variable number of setae (seven or eight) on the distal segment of the branch. (© 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Flower and fruit development in Arabidopsis thaliana

The study of flower development has experienced great advances over the last 15 years. The most important landmark was the proposal of the ABC model in which three different functions of overlapping activities account for the development of the four rings of organs of the eudicot flower. Most interestingly, during recent years this simple and elegant model has been broadly accepted and is applicable to a wide range of plant species. However, recent advances in the characterization of protein interactions and the discovery of the SEPALLATA genes that are required for proper floral organ development have led to a revision of the ABC model. The largely accepted floral quartet model, which includes the new SEPALLATA function, postulates that the development of a specific floral organ is achieved by the formation of a single complex of different MADS-box proteins. The ultimate fate of the flower is to become a fruit, ensuring dispersal of the seeds and therefore survival of the species. The Arabidopsis fruit is a silique or pod. Only in the last five years important advances have been made in establishing the differentiation of the tissues required for the opening of the fruit: the valve margins and dehiscence zone. Classical genetic analyses and molecular biology approaches have pointed to the involvement of the transcription factors SHP, ALC and IND in the formation of these tissues and of FUL and RPL in repressing this identity in the bordering tissues, valves and replum, respectively.

Prorocentrum minimum bloom and its possible link to a massive fish kill in Bolinao, Pangasinan, Northern Philippines

For the first time, a Prorocentrum minimum bloom at a maximum cell density of 4.7 × 10 5 cells/L was recorded on January 31 to February 4, 2002 at Bolinao, Pangasinan, Northern Philippines where intensive and extensive aquaculture of Chanos chanos (milkfish) in fish pens and cages has been practiced for years now. The fish kill, which lasted almost simultaneously with the bloom of the organism had its peak when the organisms bloom was declining. Lack of oxygen in the cages and pens was the fundamental cause of the fish kill. Losses due to the fish kill were estimated at six million pesos (equivalent to US$ 120,000), which includes only the worth of dead cultivated fish. Lack of oxygen in the cages and pens was the fundamental cause of the fish kill, and toxicity of the Prorocentrum could not be confirmed. The cells had minute spinules equally all over the surface of valves. Intercalary striae were wide with many ridges perpendicular to valve margin. Outline of cells was rounder than typical P. minimum cells and similar to P. balticum. Recommendations for future research on the organism are incorporated together with monitoring and management interventions in order to mitigate or possibly prevent damages in similar future events.

Chemico‐structure of the organophosphatic shells of siphonotretide brachiopods

The organophosphatic shell of siphonotretide brachiopods is stratiform with orthodoxly secreted primary and secondary layers. The dominant apatitic constituents of the secondary layer are prismatic laths and rods arranged in monolayers (occasionally in cross‐bladed successions), normally recrystallized as platy laminae. Sporadically distributed, interlaminar, lenticular chambers, containing apatitic meshes of laths and aggregates of plates and spherulites, probably represent degraded, localized exudations of glycosaminoglycans (GAGs) with dispersed apatite.The shells of Helmersenia and Gorchakovia are perforated by canals with external depressions (antechambers) that possibly contained chitinous tubercles in vivo. The immature shell of Siphonotreta and most other siphonotretids is similarly perforated and pitted; but the mature part bears recumbent, rheomorphic, hollow spines that grew forward out of pits. Internally, spines pierce the shell as independent structures to terminate as pillars in GAGs chambers. Spines and pillars were probably secreted by collectives of specialized cells (acanthoblasts) within the mantle.The shell of the oldest siphonotretide, Schizambon, is imperforate but the ventral valve has a pedicle foramen that lies forward of the posterior margin of the juvenile valve. This relationship characterizes all siphonotretides, suggesting that the pedicle, in vivo, originated within the ventral outer epithelium and not from the posterior body wall as in lingulides.

Stability Margin of Hybrid Journal Bearing: Influence of Thermal and Elastic Effects

This paper deals with the stability margin of a constant flow valve compensated hole-entry hybrid journal bearing system considering bearing flexibility and variation of viscosity due to temperature rise of the lubricant. The journal temperature is computed on the basis of average fluid-film temperature and axisymmetric isothermal element is assumed. The coupled solution of Reynold’s, energy, conduction, and elasticity equations is obtained using finite element method. The performance of a symmetric and asymmetric hole-entry hybrid journal bearing configurations is studied. The results presented in the study indicate that the bearing flexibility and temperature rise of the lubricant fluid-film affects the performance of the hole-entry hybrid journal bearing system quite significantly and proper selection of restrictor design parameter is quite useful in maintaining the fluid film thickness and threshold speed of the journal.

The alternation of different morphotypes in the seasonal cycle of the toxic diatom Pseudo-nitzschia galaxiae

The marine diatom Pseudo-nitzschia galaxiae Lundholm et Moestrup has been recently described from Mexican and Australian plankton. In this paper, we illustrate the considerable morphological variability of the species in the Mediterranean Sea and present first evidence of its toxicity. In addition to lanceolate cells 25–41 μm long, which fit the original description of the species, markedly larger (<82 μm) and smaller (>10 μm) specimens are commonly recorded. Cells of the largest size have almost parallel valve margins, while smaller specimens have extremely short rostrate ends and do not form colonies. Despite remarkable differences in shape and size, the typical ultrastructure of the species was observed for the different size classes in culture and in natural samples. In culture, cell length decreased at a rate of 1.1–2.1 μm per month. Liquid chromatography–mass spectrometry (LC–MS) analyses revealed the presence of domoic acid (DA) at very low levels in two of seven strains analyzed. LSU rDNA analysis confirmed the identity of the species and showed a very low genetic variability for the strains from the Gulf of Naples, with no relationships with size and overall shape of the cells. A relatively high number (53) of Pseudo-nitzschia sequences were considered in the phylogenetic analysis, yet the relationships among species remain unclear, probably in relation with a recent speciation process in the genus. In natural samples, P. galaxiae populations of different cell sizes occurred at different times over the year, with smaller cells found in winter and early spring, and medium and larger cells peaking in late spring–summer. The maximum concentration value in the Gulf of Naples was recorded in May 1985 (9.4×10 6 cells l −1). From the analysis of a high number of both natural and culture samples, it is concluded that size and shape variations are indicative of different stages of the life cycle of P. galaxiae, which exhibit a synchronized and seasonal occurrence at the interannual scale.