Dynamics of pigment degradation by the copepodite stage of <Emphasis Type="Italic">Pseudodiaptomus</Emphasis><Emphasis Type="Italic">euryhalinus</Emphasis> feeding on <Emphasis Type="Italic">Tetraselmis</Emphasis><Emphasis Type="Italic">suecica</Emphasis>

Short-term (3 h) changes in concentration of chlorophylls and their derivatives in stage V Pseudodiaptomus euryhalinus and their fecal material were followed by HPLC during a 24 h experiment. Copepodites were fed with the prasinophyte Tetraselmis suecica. Intact chlorophyll a and b were found in animals and fecal material and had similar dynamics of accumulation over time. The extent of transformation of chlorophyll a and b to colorless compounds was different with chlorophyll a being more extensively degraded. Additionally, several chlorophyll derivatives (pheophytin and pyropheophytin-like pigments) were found. Pyropheophytin a was the most abundant followed by pheophytin b, pheophytin a, and pheophorbide a. Relative amounts of pheopigments were different in copepodites and fecal material, and pheophytin a, pheophorbide a, and pheophytin b concentrations were low and variable. The amount of ingested chlorophyll recovered as chlorophyll a and its derivatives in fecal and copepodite pools was generally low (<5%), with one exception occurring after 9 h, when it accounted for >70%. These data suggest individual pheopigments are produced at different rates and that chemical or enzymatic mechanisms in the gut of copepodites act on the two chlorophylls in different ways.     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Viable algae released by the seastar <Emphasis Type="Italic">Dermasterias</Emphasis><Emphasis Type="Italic">imbricata</Emphasis> feeding on the symbiotic sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

Echinoderms are major predators of anemones in temperate ecosystems. The fate of two algae, zooxanthellae and zoochlorellae, after their host anemone (Anthopleura elegantissima Brandt) was consumed by the leather star Dermasterias imbricata Grube was determined in experiments conducted in July and August 2004. Productivity, photosynthetic pigments, and mitotic index (percent of cells dividing) were used as indicators of algal health; algae released after leather stars consumed their host were compared with algae freshly isolated from anemones. Two types of waste products contained algae: pellets resulting from extraoral digestion, and feces. Zooxanthellae and zoochlorellae isolated from these waste products were photosynthetic, although to different extents. For algae from feces and pellets, light-saturated photosynthetic rates (P _max) were 85 and 13%, respectively, of P _max of freshly isolated zooxanthellae; and were 20 and 46%, respectively, for zoochlorellae. The photosynthetic pigments and mitotic index (percent of dividing cells) were not altered by the feeding activities of the leather star. These results show that algae released by seastar predation on their hosts remain viable, and are hence available for establishing symbioses in A. elegantissima and other potential hosts.     

Settlement of the wreckfish (<Emphasis Type="Italic">Polyprion</Emphasis><Emphasis Type="Italic">americanus</Emphasis>)

The wreckfish (Polyprion americanus) is a panoceanic, long-lived, deep-water demersal species, characterized by an extended pelagic juvenile phase associated with floating objects. In the present study morphometric data from 435 specimens collected from Greece (Crete), Italy (Ionian Sea) and the Atlantic coast of France from September 1999 to March 2001 were analyzed to estimate the settlement size of the species and associated changes in morphology. The fishery sample included specimens from both the pelagic and the demersal stage. Length-at-settlement (TL₅₀) and 95% confidence intervals were estimated by fitting a logistic function. Eleven morphometric characters were analyzed, and the existing inflection points, reflecting alterations in body shape, were defined: (1) iteratively, as the transition point, splitting the data set into two groups, for which the reduced major axis functions, between a character and total length, of the successive groups best fit (minimum sum of squares of residuals) the combined data set and (2) where the second derivative of the fitted third-order polynomial functions to morphometric ratios equaled zero. The main size range within which wreckfish settled was 56-65 cm, and the means of the inflection points defined by the two methods were 61 and 64 cm. These results indicate changes in functional morphology associated with settlement. The monitoring of wreckfish juveniles caught in the wild and kept individually in captivity showed changes in growth and food intake at approximately the estimated settlement length and suggested that temperature was the most likely factor triggering settlement.     

Effects of substrate type on growth and mortality of blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">edulis</Emphasis>) exposed to the predator <Emphasis Type="Italic">Carcinus</Emphasis><Emphasis Type="Italic">maenas</Emphasis>

Structure and complexity of the substrate are important habitat characteristics for benthic epifauna. The specific growth and mortality rates and inducible defence characters on medium-sized blue mussels (Mytilus edulis L.) exposed to shore crabs (Carcinus maenas L.) were examined on three different substrate types in combined field and laboratory experiments. The experiments showed that complexity of the substrate increased blue mussel survival significantly, through a decrease in predation pressure. However, increased intraspecific competition for food on the complex substrate resulted in significantly lower growth rates of the mussels. Inducible defence characters were also influenced by substrate type. Blue mussels were more affected by predators on the structurally simple substrate, where they developed thicker shells and a larger posterior adductor muscle.     

Inter-polyp genetic and physiological characterisation of <Emphasis Type="Italic">Symbiodinium</Emphasis> in an <Emphasis Type="Italic">Acropora valida</Emphasis> colony

Corals harbouring genetically mixed communities of endosymbiotic algae (Symbiodinium) often show distribution patterns in accordance with differences in light climate across an individual colony. However, the physiology of these genetically characterised communities is not well understood. Single stranded conformation polymorphism (SSCP) and real time quantitative polymerase chain reaction (qPCR) analyses were used to examine the genetic diversity of the Symbiodinium community in hospite across an individual colony of Acropora valida at the spatial scale of single polyps. The physiological characteristics of the polyps were examined prior to sampling with a combined O₂ microelectrode with a fibre-optic microprobe (combined sensor diameter 50–100 μm) enabling simultaneous measurements of O₂ concentration, gross photosynthesis rate and photosystem II (PSII) quantum yield at the coral surface as a function of increasing irradiances. Both sun- and shade-adapted polyps were found to harbour either Symbiodinium clade C types alone or clades A and C simultaneously. Polyps were grouped in two categories according to (1) their orientation towardps light, or (2) their symbiont community composition. Physiological differences were not detected between sun- and shade-adapted polyps, but O₂ concentration at 1,100 μmol photons m⁻² s⁻¹ was higher in polyps that harboured both clades A and C symbionts than in polyps that harboured clade C only. These results suggest that the acclimatisation of zooxanthellae of individual polyps of an A. valida colony to ambient light levels may not be the only determinant of the photosynthetic capacity of zooxanthellae. Here, we found that photosynthetic capacity is also likely to have a strong genetic basis and differs between genetically distinct Symbiodinium types.     

Distribution of the alien ctenophore <Emphasis Type="Italic">Mnemiopsis</Emphasis><Emphasis Type="Italic">leidyi</Emphasis> in the Caspian Sea in August 2001

In this study, spatial and vertical distributions of the invasive ctenophore Mnemiopsis leidyi in the Caspian Sea were evaluated by using data collected at 41 stations during the August 2001 cruise. A comparison of data from different depths revealed that M. leidyi were generally confined to surface waters. The maximum size of the ctenophore was only 41-45 mm, and the bulk of individuals (85.5%) were <10 mm in length. The average and maximum biomasses of M. leidyi were calculated as 120 and 351 g wet weight m^-2, respectively. Whilst highest biomasses were observed in the western and central Middle Caspian Sea, hot spot areas of reproduction were present along the coasts of the western Caspian Sea, with abundance values of up to 2285 ind. m^-2. The impact of such high densities of M. leidyi is expected to be significant for the pelagic ecosystem of the Caspian Sea.     

Mate preferences in <Emphasis Type="Italic">Drosophila</Emphasis> infected with <Emphasis Type="Italic">Wolbachia</Emphasis>?

The maternally inherited bacterium Wolbachia pipientis generates strong reproductive incompatibilities between uninfected females and infected males (cytoplasmic incompatibility), significantly reducing both female and male reproductive success. Such fitness costs are thought to place selective pressure on hosts to evolve pre-copulatory preferences for mating with compatible mates, thereby enabling them to avoid the reproductive incompatibilities associated with Wolbachia. Therefore, uninfected females are predicted to prefer mating with uninfected males, whereas infected males are predicted to prefer mating with infected females. Despite these predictions, previous investigations of pre-copulatory mate preferences in Wolbachia-manipulated Drosophila have not found evidence of female preference for uninfected or compatible males. However, none of these studies utilised a design where focal individuals are provided with a simple choice in a relatively non-competitive situation. We examined both female and male pre-copulatory mate preference based on mate infection status in Drosophila simulans and D. melanogaster using simple choice assays involving between 30–50 replicates per treatment. Although we found no evidence of female pre-copulatory mate preferences in either species, male D. simulans exhibited some preference for mating with females of the same infection status. However, this preference was not evident when we repeated the experiment to confirm this finding. Consequently, we conclude that neither male nor female D. melanogaster and D. simulans exhibit significant Wolbachia-associated pre-copulatory mate preferences.     

Individual specialization in the hunting wasp <Emphasis Type="Italic">Trypoxylon</Emphasis> (<Emphasis Type="Italic">Trypargilum</Emphasis>) <Emphasis Type="Italic">albonigrum</Emphasis> (Hymenoptera,Crabronidae)

Individual-level variation in resource use occurs in a broad array of vertebrate and invertebrate taxa and may have important ecological and evolutionary implications. In this study, we measured the degree of individual-level variation in prey preference of the hunting wasp Trypoxylon albonigrum, which inhabits the Atlantic Forest in southeastern Brazil. This wasp captures several orb-weaving spider genera to provision nests. Individuals consistently specialized on a narrow subset of the prey taxa consumed by the population, indicating the existence of significant individual-level variation in prey preferences. The population niche was broader in the wet season in terms of both prey size and taxa. In the case of prey size, the population niche expansion was achieved via increased individual niche breadths, whereas in the case of prey taxa, individual niches remained relatively constrained, and the population niche expanded via increased interindividual variation. The observed pattern suggests the possibility of functional trade-offs associated with the taxon of the consumed prey. The nature of the trade-offs remains unknown, but they are likely related to learning in searching and/or handling prey. We hypothesize that by specializing on specific prey taxa, individuals increase foraging efficiency, reducing foraging time and ultimately increasing reproductive success.     

Molecular population structure of the kuruma shrimp <Emphasis Type="Italic">Penaeus</Emphasis><Emphasis Type="Italic"> japonicus</Emphasis> species complex in western Pacific

In a previous study on the kuruma shrimp Penaeus japonicus from the South China Sea, we detected high genetic divergence between two morphologically similar varieties (I and II) with distinct color banding patterns on the carapace, indicating the occurrence of cryptic species. In the present study, we clarify the geographical distribution of the two varieties in the western Pacific by investigating the genetic differentiation of the shrimp from ten localities. Two Mediterranean populations are also included for comparison. Based on the mitochondrial DNA sequence data, the shrimps are separated into two distinct clades representing the two varieties. Variety I comprises populations from Japan and China (including Taiwan), while variety II consists of populations from Southeast Asia (Vietnam, Singapore and the Philippines), Australia and the Mediterranean. Population differentiation is evident in variety II, as supported by restriction profiles of two mitochondrial markers and analysis of two microsatellite loci. The Australian population is genetically diverged from the others, whereas the Southeast Asian and Mediterranean populations show a close genetic relationship. Variety I does not occur in these three localities, while a small proportion of variety II is found along the northern coast of the South China Sea and Taiwan, which constitute the sympatric zone of the two varieties. The present study reveals high genetic diversity of P. japonicus. Further studies on the genetic structure of this species complex, particularly the populations in the Indian Ocean and Mediterranean, are needed not only to understand the evolutionary history of the shrimp, but also to improve the knowledge-based fishery management and aquaculture development programs of this important biological resource.     

Intraguild predatory interactions between the jellyfish <Emphasis Type="Italic">Cyanea capillata</Emphasis> and <Emphasis Type="Italic">Aurelia aurita</Emphasis>

While qualitative observations of jellyfish intraguild predation abound in the literature, there are only few rate measurements of these interactions. We quantified predation rates among two common jellyfish in northern boreal waters, Cyanea capillata and its prey Aurelia aurita, both of which also feed on crustacean zooplankton and fish larvae. A series of incubation experiments using a wide range of prey concentrations (0.38–3.8 m⁻³) in large containers (2.6 m³) was carried out. By replenishing the prey continuously as they were captured we maintained a nearly constant prey concentrations. Ingestion rates increased linearly up to prey concentrations of 1.92 m⁻³, yielding maximum clearance rates of ∼2.37 ± 0.39 m³ predator⁻¹ h⁻¹ for C. capillata predators 16 ± 2.3 cm in diameter. Mean ingestion rate at saturated prey concentrations (1.92–3.85 m⁻³) was 4.01 ± 0.78 prey predator⁻¹ h⁻¹. Behavioral observations suggested that predators did not alter their swimming behavior during meals, and thus that feeding rates were generally handling limited rather than encounter limited. Predators captured more prey than needed, and semi-digested prey was often discarded when fresh prey was encountered.     

Foraging responses in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae) to floral scents

Summary. For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants. Received 2 September 2001; accepted 9 September 2002.     

Antennal responses to floral scents in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae)

Summary. To better understand the biological role of floral scents for butterflies, electrophysiological responses to floral scents were investigated using combined gas chromatography and electroantennographic detection (GC-EAD). The antennal responses of three butterfly species, Aglais urticae L. (Nymphalidae), Inachis io L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae) to floral scent compounds from both natural and synthetic mixtures were examined. Floral scents were collected from the butterfly nectar plants Cirsium arvense (L.) (Asteraceae), and Buddleja davidii Franchet cv. (Loganicaeae) with dynamic head-space methods on Tenax-GR and eluted with pentane. These eluates, composed of natural floral scent blends, represent an array of compounds in their natural state. In the GC-EAD analyses eleven compounds were identified from C. arvense with the benzenoid compound phenylacetaldehyde in highest abundance. Seventeen compounds were identified from B. davidii with the irregular terpene oxoisophorone in highest abundance. Thirty-nine synthetic floral scent compounds were mixed in pentane, in equal amounts; about 35 ng were allowed to reach the antennae. The butterflies showed antennal responses to most of the floral scent compounds from both natural and synthetic blends except to the highly volatile monoterpene alkenes. Certain benzenoid compounds such as phenylacetaldehyde, monoterpenes such as linalool, and irregular terpenes such as oxoisophorone, were emitted in relatively large amounts from C. arvense and B. davidii, and elicited the strongest antennal responses. These compounds also elicited strong antennal responses when present in the synthetic scent blends. Thus, the butterflies seem to have many and /or sensitive antennal receptors for these compounds, which points to their biological importance. Moreover, these compounds are exclusively of floral scent origin. For B. davidii, which depends highly on butterflies for pollination, the exclusive floral scent compounds emitted in high abundance could be the result of an adaptive pressure to attract butterflies. Received 2 Septemter 2001; accepted 9 September 2002.     

Evidence of a spawning area of <Emphasis Type="Italic">Anguilla</Emphasis><Emphasis Type="Italic">marmorata</Emphasis> in the western North Pacific

Leptocephali of the tropical eel Anguilla marmorata have been consistently collected in the same area of the North Equatorial Current (NEC) in the western North Pacific during three consecutive cruises in June and July of 1991 (N=28) and 1994 (N=20), and July and September of 1995 (N=27), indicating that this is a spawning area of this species. These leptocephali were collected from 130°E to 142°E and 12°N to 20°N, to the west of the Mariana Islands, in 20 tows in 1991, in 13 tows in 1994 and in 17 tows in 1995, indicating a widespread presence, but a relatively low abundance. Six of these specimens (16.3-36.0 mm total length) from the 1995 cruise, which were of the typical size range of these leptocephali, were genetically confirmed to be A. marmorata in a previous study. The consistent presence of recently spawned A. marmorata leptocephali (9-20 mm) in all 3 years, suggests that the western region of the NEC is the spawning area of the northern population of A. marmorata that was identified in a recent population genetics study. These leptocephali would thus be transported westward by the NEC and then transported north into the Kuroshio Current and toward Taiwan and Japan, or south toward the southern Philippines and into the Celebes Sea by the Mindanao Current. Available evidence indicates that A. marmorata may have potentially year-round spawning, and the presence of a spawning area of this species in the same region as that of Anguilla japonica suggests that the northern population of A. marmorata has evolved a spawning migration from East Asia, the Philippines and the Celebes Sea region to the NEC area, but differs from A. japonica by having some individuals that recruit to more southern areas.     

Localization of granulatimide alkaloids in the tissues of the ascidian <Emphasis Type="Italic">Didemnum</Emphasis><Emphasis Type="Italic">granulatum</Emphasis>

Ascidians are a notable source of nitrogen-bearing secondary metabolites with a wide range of biological activities. Although many biologically active compounds have been isolated from ascidians, it is often unclear whether the animal or associated microbial symbionts such as bacteria or fungi are the true biosynthetic source of the metabolites. We have addressed the question of the biosynthetic source of the alkaloids granulatimide and isogranulatimides by localizing these compounds within the ascidian. In this work, we demonstrate that granulatimide is stored in Didemnum granulatum tunic bladder cells. Analysis by confocal fluorescence microscopy at the granulatimide emission range indicated the presence of fluorescent cells as highly vacuolated cells found dispersed in the ascidian tunic. Since this is the most exposed ascidian tissue, it is possible that these alkaloids may have a protective role, either as sunscreens and/or as feeding deterrents.     

Altered c-Fos expression demonstrates neuronal stress in mummichog, <Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>, exposed to <Emphasis Type="Italic">Pfiesteria shumwayae</Emphasis> and <Emphasis Type="Italic">Chaetoceros concavicornis</Emphasis>

To better understand sublethal effects of harmful algal blooms (HABs) on fish, mummichog, Fundulus heteroclitus (L.), were exposed in the laboratory to varying, environmentally relevant densities of Pfiesteria shumwayae (Glasgow et Burkholder, CCMP 2089, dinoflagellate) and Chaetoceros concavicornis (Mangin, CCMP 169, diatom). Two experiments were conducted during the spring of 2003 and 2004 to quantitatively examine the effects of acute (2 h) P. shumwayae and C. concavicornis algal exposure on mummichog brain activity using c-Fos expression as a marker of altered neuronal activity. Brains from HAB-exposed fish were removed, sectioned, and stained using immunocytochemistry prior to quantifying neuronal c-Fos expression. Fish exposed to P. shumwayae and C. concavicornis showed increased c-Fos expression compared to unexposed control fish. A significant dose-response relationship was observed, with increased labeling in brains of fish exposed to higher cell densities for both HAB species tested (P ≤ 0.01). Increased labeling was found in the telencephalon, optic lobes, midbrain, and portions of the medulla. The greatest increases in expression were observed in the telencephalon of P. shumwayae-exposed fish, and in the telencephalon and optic lobes of C. concavicornis-exposed fish (P ≤ 0.01). These increases in c-Fos expression are consistent with other physical and chemical stress exposures observed in fish. Neuronal stress, evidenced by c-Fos expression, demonstrates a sublethal effect of exposure and changes in brain activity in fish exposed to HAB species.     

Cloning and expression of a tauropine dehydrogenase from the marine sponge <Emphasis Type="Italic">Suberites </Emphasis><Emphasis Type="Italic">domuncula</Emphasis>

The cDNA sequence coding for tauropine dehydrogenase (TaDH) [belonging to the family of opine dehydrogenases] has been determined. Using the demosponge Suberites domuncula, we describe for the first time the tauropine dehydrogenase gene (of length 2,992 kb) from a eukaryote, consisting of two introns flanked by three exons. Moreover, two allelic variants have been identified, which are present in the different specimens either in a homozygotic or in a heterozygotic way; the data suggest an intermediary type of heritance. Phylogenetic analyses indicate that S. domuncula TaDH is only distantly related to the opine dehydrogenases from marine invertebrates; rather it comprises high sequence similarity to bacterial ornithine cyclodeaminases (OCD). In addition, expression studies revealed that the steady-state level of TaDH dropped drastically in animals, which had been exposed to elevated aeration. Antibodies raised against the recombinant sponge TaDH were used to demonstrate that S. domuncula expresses high levels of this enzyme in almost all cells. If tissue samples were kept under additional aeration no immuno-signals could be identified. A strong accumulation of the enzyme was seen around the bacteria, existing in bacteriocytes, indicating that under aerobic conditions the bacteria might produce taurine. These data suggest involvement of the sponge TaDH in the final step of the glycolytic pathway, more specifically, in regeneration of NAD(+) under anaerobic conditions. Furthermore, potential mutual influences between bacteria and sponge are discussed, claiming a horizontal gene transfer of the gene from a bacterium to the sponge. The following sequences from Suberites domuncula have been deposited (EMBL/GenBank): the cDNA encoding the tauropine [(carboxyethyl)-taurine/derived from sulfhydryl-amino acids] dehydrogenase (SD_TaDH) under AM712888; the corresponding gene under AM712889; and the fragments of the sponge-associated bacteria comprising the opine dehydrogenase domain: SUBDO_BAC1 (AM712890), SUBDO_BAC2 (AM712891), SUBDO_BAC3 (AM712892), SUBDO_BAC5 (AM712894), SUBDO_BAC6 (AM712895), SUBDO_BAC7 (AM712896), SUBDO_BAC8 (AM712897) and SUBDO_BAC9 (AM712898).