Effects of continuous and repeated dehydration on carbon fixation by bryophytes from the maritime Antarctic

 The effects of dehydration and rehydration on carbon exchange in 14 bryophytes from the maritime Antarctic were investigated using an infra-red gas analysis system. Continuous long-term (1–12 months) and repeated (1–6 one-month cycles) desiccation responses were investigated under controlled conditions. Loss of photosynthetic rate increased with length of dehydration period in all species, although some desiccation tolerance was observed even in those bryophytes from the most hydric habitats. Percentage retention of photosynthetic rate increased from hydric to xeric species, but this pattern was not repeated in terms of absolute rates of carbon fixation due to the high initial rates in the hydric species. Repeated cycles caused a greater loss of photosynthetic rate than continuous dehydration in hydric species, but the opposite situation occurred in mesic and xeric mosses. The latter groups were possibly better able to utilise the short periods of rehydration during cycles. In most bryophytes an increase in the percentage loss of photosynthetic rate following dehydration-rehydration occurred from spring to summer to autumn samples. This pattern was clearest in the hydric species and reduced in the xeric species. These variations were largely due to changes in the initial rates of photosynthesis during the growing season. It is suggested that this increased photosynthetic capacity is stress-sensitive, and is lost during either desiccation or winter freezing; the base photosynthetic capacity, being stress-tolerant, survives either of these events. The results obtained support the hypothesis that water availability is of importance in determining the distribution of bryophytes in the Antarctic. However, only the broad scale of variation in plant communities could be explained by these observations; other factors must be important in determining the finer scale of species distribution and community composition. The results are applicable to attempts to model the productivity of Antarctic bryophytes from known or predicted environmental data. Received: 15 April 1996 / Accepted: 28 September 1996     

A comparison of plasma vitamin C and E levels in two Antarctic and two temperate water fish species

Antarctic fish have a high polyunsaturated lipid content and their muscle cells have a high mitochondria density suggesting that Antarctic fish are under greater oxidative stress than temperate water fish. To test this hypothesis, the plasma concentrations of the antioxidant vitamins E and C were measured in two Antarctic fish species, Pagothenia borchgrevinki and Trematomus bernacchii, and compared with the plasma concentrations of these vitamins in two New Zealand temperate water fish species, blue cod (Parapercis colias) and banded wrasse (Notolabrus fucicola). Neither vitamin is known to be synthesised in fish and so must be obtained from the diet. The plasma from both Antarctic fish species had vitamin E concentrations five to six times higher than those found in the two temperate water fish species. However, significantly higher levels of vitamin C were only found in the plasma of T. bernacchii, a benthic Antarctic fish. The average level of vitamin C in the plasma of the cryopelagic P. borchgrevinki was approximately one-third that of T. bernacchii. The T. bernacchii plasma yielded a high range of vitamin C values, possibly reflecting differences in nutritional status among the animals captured. No beta-carotene was found in any of the fish plasma samples studied. The data suggest that even though Antarctic fish live at -1.5 degrees C they may be exposed to greater metabolic stress from free radical mediated oxidation than temperate water species.     

Open-ocean barriers to dispersal: a test case with the Antarctic Polar Front and the ribbon worm Parborlasia corrugatus (Nemertea: Lineidae)

Open-ocean environments provide few obvious barriers to the dispersal of marine organisms. Major currents and/or environmental gradients potentially impede gene flow. One system hypothesized to form an open-ocean dispersal barrier is the Antarctic Polar Front, an area characterized by marked temperature change, deep water, and the high-flow Antarctic Circumpolar current. Despite these potential isolating factors, several invertebrate species occur in both regions, including the broadcast-spawning nemertean worm Parborlasia corrugatus. To empirically test for the presence of an open-ocean dispersal barrier, we sampled P. corrugatus and other nemerteans from southern South America, Antarctica, and the sub-Antarctic islands. Diversity was assessed by analyzing mitochondrial 16S rRNA and cytochrome c oxidase subunit I sequence data with Bayesian inference and tcs haplotype network analysis. Appropriate neutrality tests were also employed. Although our results indicate a single well-mixed lineage in Antarctica and the sub-Antarctic, no evidence for recent gene flow was detected between this population and South American P. corrugatus. Thus, even though P. corrugatus can disperse over large geographical distances, physical oceanographic barriers (i.e. Antarctic Polar Front and Antarctic Circumpolar Current) between continents have likely restricted dispersal over evolutionary time. Genetic distances and haplotype network analysis between South American and Antarctic/sub-Antarctic P. corrugatus suggest that these two populations are possibly two cryptic species.     

Biodiversity and biogeography of Southern Ocean pycnogonids

The pycnogonids of the Southern Ocean have been studied for almost two centuries and have played a key role in shaping previous biogeographic regions for the Antarctic benthos. The aim of this study was to assess the biogeographic patterns derived from the most current sample records of pycnogonids from the Southern Ocean and neighbouring areas. 332 species of pycnogonids from 1837 sample locations were analysed using 279 3° by 3° grid cells. We investigated richness patterns and the effect of sampling intensity at both local and regional scales, and used multivariate analysis of distribution patterns and species assemblages to define biogeographic trends. These analyses identified a distinct and isolated Antarctic pycnogonid shelf fauna which was different to that of the deep‐sea around Antarctica, the Sub‐Antarctic islands, South America or New Zealand. Within the Antarctic, we found the South Shetland Islands to be the most speciose region and a probable center of radiation for the pycnogonids. No latitudinal gradients in species richness were detected. We note that the distribution patterns observed are based upon classical taxonomy and discuss the potential for changes to these patterns with new insights from molecular techniques. We conclude that, even with the potential for cryptic species, the large‐scale biogeographic trends observed in the pycnogonids should hold true.     

Are Antarctic minke whales unusually abundant because of 20th century whaling?

Severe declines in megafauna worldwide illuminate the role of top predators in ecosystem structure. In the Antarctic, the Krill Surplus Hypothesis posits that the killing of more than 2 million large whales led to competitive release for smaller krill‐eating species like the Antarctic minke whale. If true, the current size of the Antarctic minke whale population may be unusually high as an indirect result of whaling. Here, we estimate the long‐term population size of the Antarctic minke whale prior to whaling by sequencing 11 nuclear genetic markers from 52 modern samples purchased in Japanese meat markets. We use coalescent simulations to explore the potential influence of population substructure and find that even though our samples are drawn from a limited geographic area, our estimate reflects ocean‐wide genetic diversity. Using Bayesian estimates of the mutation rate and coalescent‐based analyses of genetic diversity across loci, we calculate the long‐term population size of the Antarctic minke whale to be 670 000 individuals (95% confidence interval: 374 000–1 150 000). Our estimate of long‐term abundance is similar to, or greater than, contemporary abundance estimates, suggesting that managing Antarctic ecosystems under the assumption that Antarctic minke whales are unusually abundant is not warranted.     

Antarctic Crabs: Invasion or Endurance?

Recent scientific interest following the “discovery” of lithodid crabs around Antarctica has centred on a hypothesis that these crabs might be poised to invade the Antarctic shelf if the recent warming trend continues, potentially decimating its native fauna. This “invasion hypothesis” suggests that decapod crabs were driven out of Antarctica 40–15 million years ago and are only now returning as “warm” enough habitats become available. The hypothesis is based on a geographically and spatially poor fossil record of a different group of crabs (Brachyura), and examination of relatively few Recent lithodid samples from the Antarctic slope. In this paper, we examine the existing lithodid fossil record and present the distribution and biogeographic patterns derived from over 16,000 records of Recent Southern Hemisphere crabs and lobsters. Globally, the lithodid fossil record consists of only two known specimens, neither of which comes from the Antarctic. Recent records show that 22 species of crabs and lobsters have been reported from the Southern Ocean, with 12 species found south of 60°S. All are restricted to waters warmer than 0°C, with their Antarctic distribution limited to the areas of seafloor dominated by Circumpolar Deep Water (CDW). Currently, CDW extends further and shallower onto the West Antarctic shelf than the known distribution ranges of most lithodid species examined. Geological evidence suggests that West Antarctic shelf could have been available for colonisation during the last 9,000 years. Distribution patterns, species richness, and levels of endemism all suggest that, rather than becoming extinct and recently re-invading from outside Antarctica, the lithodid crabs have likely persisted, and even radiated, on or near to Antarctic slope. We conclude there is no evidence for a modern-day “crab invasion”. We recommend a repeated targeted lithodid sampling program along the West Antarctic shelf to fully test the validity of the “invasion hypothesis”.     

Use of chemical tracers to assess diet and persistent organic pollutants in Antarctic Type C killer whales

Measuring chemical tracers in tissues of marine predators provides insight into the prey consumed and the predator's contaminant exposure. In this study, samples from Type C killer whales ( Orcinus orca ) biopsied in Antarctica were analyzed for chemical tracers ( i.e. , stable isotopes of carbon and nitrogen, fatty acids, and persistent organic pollutants [POPs]). Profiles of these individual tracers were very different from those of killer whale populations that have been studied in the eastern North and eastern Tropical Pacific. For example, δ¹³C and δ¹⁵N stable isotope values and most POP concentrations were significantly lower in the Antarctic population. In addition, multivariate statistical analyses of both fatty acid and POP profiles found distinctly different patterns for Antarctic Type C whales compared to those from whales in the other populations. Similar assays were conducted on four species of Antarctic marine fish considered potential prey for Type C killer whales. Results were consistent with a diet of fish for Type C whales, but other species ( e.g. , low trophic-level marine mammals or penguins) could not be eliminated as supplemental prey.     

Isolation, characterization, and ecology of cold-active, chemolithotrophic, sulfur-oxidizing bacteria from perennially ice-covered Lake Fryxell, Antarctica

Novel strains of obligately chemolithoautotrophic, sulfur-oxidizing bacteria have been isolated from various depths of Lake Fryxell, Antarctica. Physiological, morphological, and phylogenetic analyses showed these strains to be related to mesophilic Thiobacillus species, such as T. thioparus. However, the psychrotolerant Antarctic isolates showed an adaptation to cold temperatures and thus should be active in the nearly freezing waters of the lake. Enumeration by most-probable-number analysis in an oxic, thiosulfate-containing medium revealed that the sulfur-oxidizing chemolithotroph population peaks precisely at the oxycline (9.5 m), although viable cells exist well into the anoxic, sulfidic waters of the lake. The sulfur-oxidizing bacteria described here likely play a key role in the biogeochemical cycling of carbon and sulfur in Lake Fryxell.     

Pseudomonas, the dominant polycyclic aromatic hydrocarbon-degrading bacteria isolated from Antarctic soils and the role of large plasmids in horizontal gene transfer

Twenty-two polycyclic aromatic hydrocarbon (PAH)-degrading bacterial strains were isolated from Antarctic soils with naphthalene or phenanthrene as a sole carbon source, while no degrader was obtained from an unpolluted sampling site. Phylogenetic analysis showed that all belonged to the genus Pseudomonas except one that was identified as the genus of Rahnella. Some of them were closely related to previously reported cold-tolerant species, while some were separated in deeply rooted branches and represent new strains. All these strains showed a high efficiency to degrade naphthalene at 4 degrees C, and some additionally degraded phenanthrene. Using degenerate primers and polymerase chain reaction (PCR) amplification, ndo gene encoding naphthalene dioxygenase (NDO) was detected from all the isolates. Phylogenetic analysis grouped these genes into two clusters which shared 94% similarity to each other, and showed about 97% similarity within a cluster. However, no obvious difference was observed with mesophilic ndo genes; this indicates that the host cell is pivotal in cold adaptation. In addition, the mismatch between 16S rRNA and NDO phylogenetic trees strongly indicates horizontal gene transfer among these isolates and may have happened in situ. Further, Southern hybridization and plasmid curing confirmed that ndo genes were located on a large self-transmissible plasmid, which can be transferred to a mesophilic strains. The transconjugants acquired the ability to utilize naphthalene and phenanthrene. Results of this article imply that Pseudomonas plays an important role in PAH biodegradation in Antarctic soils, and the related genes might be originally transferred from outside Antarctica and spread among indigenous species.     

Fungi in Antarctica

Fungi are generally easily dispersed and are able to colonize a very wide variety of different substrata and to withstand many different environmental conditions. Because of these characteristics they spread all over the world. The Antarctic mycoflora is quite diversified within the different climatic regions of the continent. Most Antarctic microfungi are cosmopolitan; some of them are propagules transported to Antarctica but unable to grow under the Antarctic conditions, while others, termed indigenous, are well adapted and able to grow and reproduce even at low temperatures, mostly as psychrotolerant, or fast sporulating forms, able to conclude their life-cycles in very short time. In the most extreme and isolated areas of the continent, such as the Antarctic Dry Valleys, endemic species showing physiological and morphological adaptations have locally evolved. Most Antarctic fungi, as well as fungi from other dry and cold habitats, are adapted to low temperatures, repeated freeze and thawing cycles, low water availability, osmotic stress, desiccation, low nutrients availability and high UV radiation. Sometimes single strategies are not specific for single stress factors and allow these microorganisms to cope with more than one unfavourable condition.     

Do bipolar distributions exist in marine sponges? Stylocordyla chupachups sp. nv. (Porifera: Hadromerida) from the Weddell Sea (Antarctic), previously reported as S. borealis (Lov��n, 1868)

Sponges are the dominant invertebrates in many Antarctic communities, where they play a decisive structural function thanks to their abundance and large sizes. However, current knowledge on Antarctic sponges remains poor even with respect to basic aspects such as taxonomy. Here, we report on an Antarctic species of the genus Stylocordyla, which has been recorded for a long time under the name of the boreal S. borealis due to spicule and growth habit similarities. A thorough study of dense populations of the only Stylocordyla species known up to now from the eastern zone of the Weddell Sea as well as the re-examination of several specimens (including the type material) of S. borealis has allowed us to assess the variability of the boreal species and to confirm that the austral species is not S. borealis (Lovén, 1868) but a new species of Stylocordyla, different from the other congeners recorded from southern latitudes. The new species S. chupachups commonly dwells on horizontal or slighted sloped hard bottoms of the continental shelf of Weddell Sea, from 100 m to below 400 m depth, although the densest populations usually occur between 150 and 300 m. It is a pioneer species in areas that have been scoured by icebergs, and thus its presence may be considered an indicator of recent colonisation.     

Mega-epibenthos at Bouvet Island (South Atlantic): a spatially isolated biodiversity hot spot on a tiny geological spot

Mega-epibenthic diversity was analysed using a seabed photography at four stations off Bouvet Island and one station at the Spiess Seamount in the South Atlantic. Surprisingly, the intermediate-scale diversity within the area of investigation was not lower compared to that on the Patagonian shelf and only moderately lower than that on the Antarctic continental shelf. This result is incompatible with Mac Arthur and Wilson’s Island Biogeography Theory describing species richness as a function of immigration of new species into an area and its extension. The relatively high species number and the very small extension of the Bouvet shelf compared to the much larger continental shelves of the other two areas can be explained by long-range dispersal of marine benthic animals in the Antarctic Circumpolar Current and high habitat heterogeneity. The observed uncoupling of intermediate-scale from large-scale background species diversity on the Antarctic shelf raises the question whether in these benthic systems an upper capacity limit for diversity exists.     

Hydrocarbons in the seawater and pelagic organisms of the Southern Ocean

Summary The aliphatic and aromatic hydrocarbon content of seawater and a selection of marine pelagic organisms from the Bransfield Strait, Antarctica were evaluated. The patterns of the hydrocarbons found indicated that their origin was biogenic and there was no evidence for anthropogenic hydrocarbons in the Antarctic marine ecosystem. Hydrocarbons which are constituents of man made materials are present throughout the region but not in the patterns characteristic of anthropogenic origin. Recent reports of an even carbon number predominance in Antarctic marine organisms were not corroborated in this study. A number of biogenic compounds were investigated as suitable markers to monitor pathways of hydrocarbons in the Antarctic food chain. Specific compounds (probably branched chain alkenes) were found to occur at more than one trophic level. Polyaromatic hydrocarbons were present in all organisms but were undetectable in 44% of the seawater samples.     

Evolution and biodiversity of Antarctic organisms: a molecular perspective

The Antarctic biota is highly endemic, and the diversity and abundance of taxonomic groups differ from elsewhere in the world. Such characteristics have resulted from evolution in isolation in an increasingly extreme environment over the last 100 Myr. Studies on Antarctic species represent some of the best examples of natural selection at the molecular, structural and physiological levels. Analyses of molecular genetics data are consistent with the diversity and distribution of marine and terrestrial taxa having been strongly influenced by geological and climatic cooling events over the last 70 Myr. Such events have resulted in vicariance driven by continental drift and thermal isolation of the Antarctic, and in pulses of species range contraction into refugia and subsequent expansion and secondary contact of genetically distinct populations or sister species during cycles of glaciation. Limited habitat availability has played a major role in structuring populations of species both in the past and in the present day. For these reasons, despite the apparent simplicity or homogeneity of Antarctic terrestrial and marine environments, populations of species are often geographically structured into genetically distinct lineages. In some cases, genetic studies have revealed that species defined by morphological characters are complexes of cryptic or sibling species. Climate change will cause changes in the distribution of many Antarctic and sub-Antarctic species through affecting population-level processes such as life history and dispersal.     

Relationship between the heat tolerance of photosynthesis and the thermal stability of rubisco activase in plants from contrasting thermal environments

Inhibition of net photosynthesis (Pn) by moderate heat stress has been attributed to an inability of Rubisco activase to maintain Rubisco in an active form. To examine this proposal, the temperature response of Pn, Rubisco activation, chlorophyll fluorescence, and the activities of Rubisco and Rubisco activase were examined in species from contrasting environments. The temperature optimum of Rubisco activation was 10 degrees C higher in the creosote bush (Larrea tridentata) compared with the Antarctic hairgrass (Deschampsia antarctica), resembling the temperature response of Pn. Pn increased markedly with increasing internal CO(2) concentration in Antarctic hairgrass and creosote bush plants subjected to moderate heat stress even under nonphotorespiratory conditions. Nonphotochemical quenching of chlorophyll fluorescence, the effective quantum yield of photochemical energy conversion (DeltaF/F(m)') and the maximum yield of PSII (F(v)/F(m)) were more sensitive to temperature in Antarctic hairgrass and two other species endemic to cold regions (i.e. Lysipomia pumila and spinach [Spinacea oleracea]) compared with creosote bush and three species (i.e. jojoba [Simmondsia chinensis], tobacco [Nicotiana tabacum], and cotton [Gossypium hirsutum]) from warm regions. The temperature response of activity and the rate of catalytic inactivation of Rubisco from creosote bush and Antarctic hairgrass were similar, whereas the optimum for ATP hydrolysis and Rubisco activation by recombinant creosote bush, cotton, and tobacco activase was 8 degrees C to 10 degrees C higher than for Antarctic hairgrass and spinach activase. These results support a role for activase in limiting photosynthesis at high temperature.     

Evidence of hidden biodiversity, ongoing speciation and diverse patterns of genetic structure in giant Antarctic amphipods

Recent molecular research on Antarctic benthic organisms has challenged traditional taxonomic classifications, suggesting that our current perceptions of Antarctic biodiversity and species distributions must be thoroughly revised. Furthermore, genetic differentiation at the intraspecific level remains poorly understood, particularly in eastern Antarctica. We addressed these issues using DNA sequence data for two sibling amphipod species that could be collected on a circum-Antarctic scale: Eusirus perdentatus and Eusirus giganteus. Haplotype networks and Bayesian phylogenies based on mitochondrial (COI, CytB) and nuclear (ITS2) DNA provided strong evidence of multiple cryptic species of Eusirus, with several occurring in sympatry and at least one likely to have a true circum-Antarctic distribution. Within species, gene flow was often highly restricted, consistent with a brooding life history and in some cases suggestive of current or future allopatric speciation. Patterns of genetic structure were not always predictable: one cryptic species showed preliminary evidence of high genetic differentiation across ～150 km in eastern Antarctica (F(ST) > 0.47, P < 0.01), yet another was remarkably homogenous across ～5000 km (F(ST) = 0.00, P = 1.00). Genetic diversity also varied among cryptic species, independent of sample size (π = 0.00-0.99). These results indicate several hidden levels of genetic complexity in these Antarctic amphipods that are neither apparent from previous taxonomic or ecological studies nor predictable from their life history. Such genetic diversity and structure may reflect different modes of survival for Antarctic benthic organisms during historic glacial cycles, and/or subsequent re-establishment of populations on the shelf, and highlight our misunderstanding of Antarctic marine species diversity.     

Ecophysiology of Antarctic vascular plants

Most of the ice and snow-free land in the Antarctic summer is found along the Antarctic Peninsula and adjacent islands and coastal areas of the continent. This is the area where most of the Antarctic vegetation is found. Mean air temperature tends to be above zero during the summer in parts of the Maritime Antarctic. The most commonly found photosynthetic organisms in the Maritime Antarctic and continental edge are lichens (around 380 species) and bryophytes (130 species). Only two vascular plants, Deschampsia antarctica Desv. and Colobanthus quitensis (Kunth) Bartl., have been able to colonize some of the coastal areas. This low species diversity, compared with the Arctic, may be due to permanent low temperature and isolation from continental sources of propagules. The existence of these plants in such a permanent harsh environment makes them of particular interest for the study of adaptations to cold environments and mechanisms of cold resistance in plants. Among these adaptations are high freezing resistance, high resistance to light stress and high photosynthetic capacity at low temperature. In this paper, the ecophysiology of the two vascular plants is reviewed, including habitat characteristics, photosynthetic properties, cold resistance, and biochemical adaptations to cold.     

Photosynthesis,dark respiration and light independent carbon fixation of endemic Antarctic macroalgae

Summary The light saturated photosynthesis, dark respiration and light independent carbon fixation of macroalgal species endemic to the Antarctic were measured. Five brown algae. Ascoseira mirabilis, Desmarestia anceps, D. antarctica, Phaeurus antarcticus, Himantothallus grandifolius and the red alga Palmaria decipiens were included. Rates of these three parameters at 0°C were very similar to those measured in other studies on temperate algae at higher temperature. This indicates a high degree of physiological adaptation to the Antarctic environment within these species. A comparison was made of polarographic and chemical means of measuring oxygen flux during photosynthesis and dark respiration at low temperature. There was a good correlation between measurements of oxygen evolution and carbon fixation, although apparent photosynthetic quotient values were in most cases high.Abbreviations HEPES N-(Hydroxyethyl) piperazine-N-(2 ethane-sulphonic acid) - RuBP D-ribulose 1,5-bisphosphate Contribution No. 415 from the Alfred-Wegener-Institut für Polaru. Meeresforschung     

Potential Climate Change Effects on the Habitat of Antarctic Krill in the Weddell Quadrant of the Southern Ocean

Antarctic krill is a cold water species, an increasingly important fishery resource and a major prey item for many fish, birds and mammals in the Southern Ocean. The fishery and the summer foraging sites of many of these predators are concentrated between 0° and 90°W. Parts of this quadrant have experienced recent localised sea surface warming of up to 0.2°C per decade, and projections suggest that further widespread warming of 0.27° to 1.08°C will occur by the late 21^st century. We assessed the potential influence of this projected warming on Antarctic krill habitat with a statistical model that links growth to temperature and chlorophyll concentration. The results divide the quadrant into two zones: a band around the Antarctic Circumpolar Current in which habitat quality is particularly vulnerable to warming, and a southern area which is relatively insensitive. Our analysis suggests that the direct effects of warming could reduce the area of growth habitat by up to 20%. The reduction in growth habitat within the range of predators, such as Antarctic fur seals, that forage from breeding sites on South Georgia could be up to 55%, and the habitat’s ability to support Antarctic krill biomass production within this range could be reduced by up to 68%. Sensitivity analysis suggests that the effects of a 50% change in summer chlorophyll concentration could be more significant than the direct effects of warming. A reduction in primary production could lead to further habitat degradation but, even if chlorophyll increased by 50%, projected warming would still cause some degradation of the habitat accessible to predators. While there is considerable uncertainty in these projections, they suggest that future climate change could have a significant negative effect on Antarctic krill growth habitat and, consequently, on Southern Ocean biodiversity and ecosystem services.