COLONIZATION AND GROWTH OF ASCOPHYLLUM NODOSUM (PHAEOPHYTA) IN MAINE1

Colonization and growth of Ascophyllum nodosum (L.) Le Jol. were investigated in two Maine estuaries from 1972 to 1978. Following denudation of intertidal rock, substrata were initially colonized by Fucus vesiculosus L.; eventually, Ascophyllum supplanted Fucus, and became dominant in terms of percentage cover. Ascophyllum settled first and most densely in the low intertidal zone, but its fastest growth occurred in the mid-intertidal zone. Some, but not all, Ascophyllum germlings produced a vesicle within one year of colonization. The mean annual growth of A. nodosum was variable among sites, zones and years.     

Seasonal growth and reproduction of estuarine fucoid algae in New England

Estuarine populations of Ascophyllum nodosum (L.) Le Jolis and Fucus vesiculosus L. var. spiralis Farlow from the Piscataqua River of the Great Bay Estuary System (New Hampshire-Maine) showed their maximum growth during the spring and fall of two consecutive years. The maximum growth rates (in terms of elongation) for Ascophyllum and Fucus were 3.5 and 3.6 cm/month, respectively, during 1973, and 3.2 and 2.3 cm/month in 1974. The average length of whole Fucus fronds showed a trend towards longer plants during the early spring and late summer-fall, while the average weight was significantly greater in the spring and early summer. Ascophyllum showed a distinct reproductive periodicity, with a maximum in March or April of both years; after the spring period of maximum reproduction, the receptacles dehisced simultaneously and the plants were vegetative during the summer. The reproductive periodicity of Fucus was not so distinct as that of Ascophyllum; even so, its maximum reproduction was in April of both years. The level of reproduction for Ascophyllum populations was three times greater in 1974 than in 1973. The reproduction and growth of the two fucoid algae are related to seasonal variations in temperature and light.     

PRESENCE OF ALGAE IN FRESHWATER ICE COVER OF FLUVIAL LAC SAINT‐PIERRE (ST. LAWRENCE RIVER,CANADA)1

Winter ice cover is a fundamental feature of north temperate aquatic systems and is associated with the least productive months of the year. Here we describe a previously unknown freshwater habitat for algal and microbial communities in the ice cover of the freshwater St. Lawrence River, Quebec, Canada. Sampling performed during winter 2005 revealed the presence of viable algal cells, such as Aulacoseira islandica (O. Müll.) Simonsen (Bacillariophyceae), and microbial assemblage growing in the ice and at the ice–water interface. Vertical channels (1–5 mm wide) containing algae were also observed. Concentrations of chl a ranged between 0.5 and 169 μg · L^?1 of melted ice, with maximal concentrations found in the lower part of the ice cores. These algae have the potential to survive when ice breakup occurs and reproduce rapidly in spring/summer conditions. Freshwater ice algae can thus contribute to in situ primary production, biodiversity, and annual carbon budget in various habitats of riverine communities.     

Use of the intertidal zone by fish in Nova Scotia

Synopsis In Nova Scotia the annual harvest of the intertidal seaweed Ascophyllum nodosum exceeds 20 000 tons. This study investigated the possible impact of removal of the seaweed on intertidal abundance of fishes and their gut contents. At sites near Lower Argyle, Nova Scotia, absolute abundance of fish in small areas with Ascophyllum removed and intact was determined with pop-up seines. Gut contents were investigated from fish collected with the seines and with trammel nets. Number and weight of fish were not significantly different between cleared and intact areas and averaged 4.6 individuals and 160 g per 400 m² during June to October. Cunner and sculpins had more food in their guts when leaving than when entering the intertidal, but only cunner had more food in their stomachs when leaving intact areas than when leaving cleared areas. Most fish captured in the intertidal were small (< 15 cm) and of no commercial value, and the numbers of commercial species were too low to be of much value. Numbers of fish in the intertidal was significantly lower than in the shallow subtidal. This study provided no evidence for adverse effects of the removal of patches of Ascophyllum on fishes.     

Persistence of bacterial and archaeal communities in sea ice through an Arctic winter

The structure of bacterial communities in first‐year spring and summer sea ice differs from that in source seawaters, suggesting selection during ice formation in autumn or taxon‐specific mortality in the ice during winter. We tested these hypotheses by weekly sampling (January–March 2004) of first‐year winter sea ice (Franklin Bay, Western Arctic) that experienced temperatures from ?9°C to ?26°C, generating community fingerprints and clone libraries for Bacteria and Archaea. Despite severe conditions and significant decreases in microbial abundance, no significant changes in richness or community structure were detected in the ice. Communities of Bacteria and Archaea in the ice, as in under‐ice seawater, were dominated by SAR11 clade Alphaproteobacteria and Marine Group I Crenarchaeota, neither of which is known from later season sea ice. The bacterial ice library contained clones of Gammaproteobacteria from oligotrophic seawater clades (e.g. OM60, OM182) but no clones from gammaproteobacterial genera commonly detected in later season sea ice by similar methods (e.g. Colwellia, Psychrobacter). The only common sea ice bacterial genus detected in winter ice was Polaribacter. Overall, selection during ice formation and mortality during winter appear to play minor roles in the process of microbial succession that leads to distinctive spring and summer sea ice communities.     

Drivers of apoplastic freezing in gymnosperm and angiosperm branches

It is not well understood what determines the degree of supercooling of apoplastic sap in trees, although it determines the number and duration of annual freeze–thaw cycles in a given environment. We studied the linkage between apoplastic ice nucleation temperature, tree water status, and conduit size. We used branches of 10 gymnosperms and 16 angiosperms collected from an arboretum in Helsinki (Finland) in winter and spring. Branches with lower relative water content froze at lower temperatures, and branch water content was lower in winter than in spring. A bench drying experiment with Picea abies confirmed that decreasing branch water potential decreases apoplastic ice nucleation temperature. The studied angiosperms froze on average 2.0 and 1.8°C closer to zero Celsius than the studied gymnosperms during winter and spring, respectively. This was caused by higher relative water content in angiosperms; when branches were saturated with water, apoplastic ice nucleation temperature of gymnosperms increased to slightly higher temperature than that of angiosperms. Apoplastic ice nucleation temperature in sampled branches was positively correlated with xylem conduit diameter as shown before, but saturating the branches removed the correlation. Decrease in ice nucleation temperature decreased the duration of freezing, which could have an effect on winter embolism formation via the time available for gas escape during ice propagation. The apoplastic ice nucleation temperature varied not only between branches but also within a branch between consecutive freeze–thaw cycles demonstrating the stochastic nature of ice nucleation.     

Multiple effects of precipitation onSalix lasiolepis and populations of the stem-galling sawfly,Euura lasiolepis

Relatively low winter precipitation (e.g., 18–28 cm from October to May compared to 45 to 65 cm) caused reduced growth of the arroyo willow,Salix lasiolepis, with number of shoots per stem initiated and shoot length reduced. Resources were reduced for the stem-galling sawfly,Euura lasiolepis, which declined in numbers after the relatively dry winter of 1980–81. Sawfly phenology was advanced relative to willow phenology in the 1981 generation, causing an additional reduction in resource availability. These direct effects of precipitation on the sawfly were increased by indirect effects on survivorship of the 1981 generation. Egg death in galls increased on water-stressed willow plants, forming the major mortality in the generation. This resulted in very low survival in the generation and an even lower population in the 1982 generation, even though resources recovered after high precipitation during the 1981–82 winter (46.7 cm). An experiment using high, medium, and low water treatments on potted willow plants demonstrated that the effects on willows and sawflies could be reproduced using only water as a variable.     

Dependency of Antarctic zooplankton species on ice algae‐produced carbon suggests a sea ice‐driven pelagic ecosystem during winter

How the abundant pelagic life of the Southern Ocean survives winter darkness, when the sea is covered by pack ice and phytoplankton production is nearly zero, is poorly understood. Ice‐associated (“sympagic”) microalgae could serve as a high‐quality carbon source during winter, but their significance in the food web is so far unquantified. To better understand the importance of ice algae‐produced carbon for the overwintering of Antarctic organisms, we investigated fatty acid (FA) and stable isotope compositions of 10 zooplankton species, and their potential sympagic and pelagic carbon sources. FA‐specific carbon stable isotope compositions were used in stable isotope mixing models to quantify the contribution of ice algae‐produced carbon (α_Ice) to the body carbon of each species. Mean α_Ice estimates ranged from 4% to 67%, with large variations between species and depending on the FA used for the modelling. Integrating the α_Ice estimates from all models, the sympagic amphipod Eusirus laticarpus was the most dependent on ice algal carbon (α_Ice: 54%–67%), and the salp Salpa thompsoni showed the least dependency on ice algal carbon (α_Ice: 8%–40%). Differences in α_Ice estimates between FAs associated with short‐term vs. long‐term lipid pools suggested an increasing importance of ice algal carbon for many species as the winter season progressed. In the abundant winter‐active copepod Calanus propinquus, mean α_Ice reached more than 50% in late winter. The trophic carbon flux from ice algae into this copepod was between 3 and 5 mg C m^?2 day^?1. This indicates that copepods and other ice‐dependent zooplankton species transfer significant amounts of carbon from ice algae into the pelagic system, where it fuels the food web, the biological carbon pump and elemental cycling. Understanding the role of ice algae‐produced carbon in these processes will be the key to predictions of the impact of future sea ice decline on Antarctic ecosystem functioning.     

Impact of early and late winter icing events on sub‐arctic dwarf shrubs

Polar regions are predicted to undergo large increases in winter temperature and an increased frequency of freeze–thaw cycles, which can cause ice layers in the snow pack and ice encasement of vegetation. Early or late winter timing of ice encasement could, however, modify the extent of damage caused to plants. To determine impacts of the date of ice encasement, a novel field experiment was established in sub‐arctic Sweden, with icing events simulated in January and March 2008 and 2009. In the subsequent summers, reproduction, phenology, growth and mortality, as well as physiological indicators of leaf damage were measured in the three dominant dwarf shrubs: Vaccinium uliginosum, Vaccinium vitis‐idaea and Empetrum nigrum. It was hypothesised that January icing would be more damaging compared to March icing due to the longer duration of ice encasement. Following 2 years of icing, E. nigrum berry production was 83% lower in January‐iced plots compared to controls, and V. vitis‐idaea electrolyte leakage was increased by 69%. Conversely, electrolyte leakage of E. nigrum was 25% lower and leaf emergence of V. vitis‐idaea commenced 11 days earlier in March‐iced plots compared to control plots in 2009. There was no effect of icing on any of the other parameters measured, indicating that overall these study species have moderate to high tolerance to ice encasement. Even much longer exposure under the January icing treatment does not clearly increase damage.     

Winter warming as an important co‐driver for Betula nana growth in western Greenland during the past century

Growing season conditions are widely recognized as the main driver for tundra shrub radial growth, but the effects of winter warming and snow remain an open question. Here, we present a more than 100 years long Betula nana ring‐width chronology from Disko Island in western Greenland that demonstrates a highly significant and positive growth response to both summer and winter air temperatures during the past century. The importance of winter temperatures for Betula nana growth is especially pronounced during the periods from 1910–1930 to 1990–2011 that were dominated by significant winter warming. To explain the strong winter importance on growth, we assessed the importance of different environmental factors using site‐specific measurements from 1991 to 2011 of soil temperatures, sea ice coverage, precipitation and snow depths. The results show a strong positive growth response to the amount of thawing and growing degree‐days as well as to winter and spring soil temperatures. In addition to these direct effects, a strong negative growth response to sea ice extent was identified, indicating a possible link between local sea ice conditions, local climate variations and Betula nana growth rates. Data also reveal a clear shift within the last 20 years from a period with thick snow depths (1991–1996) and a positive effect on Betula nana radial growth, to a period (1997–2011) with generally very shallow snow depths and no significant growth response towards snow. During this period, winter and spring soil temperatures have increased significantly suggesting that the most recent increase in Betula nana radial growth is primarily triggered by warmer winter and spring air temperatures causing earlier snowmelt that allows the soils to drain and warm quicker. The presented results may help to explain the recently observed ‘greening of the Arctic’ which may further accelerate in future years due to both direct and indirect effects of winter warming.     

Mitochondrial Activity and Ethanol Accumulation in Ice-encased Winter Cereal Seedlings

Cold-hardened dark-grown seedlings of winter wheat (Triticum aestivum L.) and winter rye (Secale cereale L.) are killed during total encasement in ice at −1 C at a rate related to the initial cold hardiness of the cultivars. Few plants remain alive after 7 days of encasement. Nonhardened seedlings are rapidly killed in ice. The respiratory properties of mitochondria isolated from plants after increasing periods of ice encasement decline slowly, and activity is little impaired when intact plants are about 50% killed. Electron microscopy indicates that mitochondrial structure is not disrupted until 3 weeks of ice encasement. Ethanol accumulates in hardened and nonhardened plants in ice, but at levels which are not toxic to the plants.     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Maintenance of the supercooled state in the gut fluid of overwintering pyrochroid beetle larvae, Dendroides canadensis : role of ice nucleators and antifreeze proteins

  The effect of gut fluid ice nucleators and antifreeze proteins on maintenance of supercooling was explored in fire-colored beetle larvae, Dendroides canadensis, via seasonal monitoring of supercooling points, antifreeze protein activity and ice nucleator activity of gut fluid and/or larvae. During cold hardening in the field, freeze-avoiding larvae evacuated their guts and depressed larval supercooling points. Analysis of gut fluid indicated supercooling points and ice nucleator activity decreased, whereas antifreeze protein activity increased as winter approached. Suspensions of bacteria isolated from guts of feeding larvae collected in spring/summer had higher supercooling points than those from midwinter-collected non-feeding larvae, suggesting bacterial ice nucleators are removed from midwinter gut fluid. The ice nucleation active bacterium Pseudomonas fluorescens was isolated from gut fluid of feeding larvae but was absent in winter. When mixed with purified D.␣canadensis hemolymph antifreeze proteins (structurally similar and/or identical to those in gut fluid), the cumulative ice nucleus spectra of P. fluorescens suspensions were shifted to lower temperatures indicating an inhibitory effect on the bacteria's ice-nucleating phenotype. By extending larval supercooling capacity, both gut clearing and masking of bacterial ice nucleators by antifreeze proteins may contribute to overwintering survival in supercooled insects. Accepted: 8 August 1996     

Metabolic acclimation to hypoxia in winter cereals : low temperature flooding increases adenylates and survival in ice encasement

Cold hardened seedlings of winter wheat (Triticum aestivum L. em Thell) show an hypoxic hardening response: an exposure to low temperature flooding increases the tolerance of plants to a subsequent ice encasement exposure. Seedlings of winter barley (Hordeum vulgare L.) do not show such a response in similar experimental conditions. During ice encasement, there are general declines in adenylate energy charge (AEC), total adenylates and ATP:ADP ratios in the crown tissues of two winter wheat cultivars, and a winter barley, but rates of decline are faster in the barley. When the ice period is preceded by low temperature flooding of the whole plant, levels of the adenylate components are raised significantly in the wheats, and to a lesser extent in the barley. The survival of plants in ice preceded by flooding is related to the increased initial level of adenylates at the onset of the ice encasement stress, and the maintenance of higher levels of adenylates and ATP in the early stages of ice encasement as a result of accelerated rates of glycolysis. Higher survival of both winter wheat and barley plants during ice encasement in the light is also associated with significantly higher levels of AEC and adenylates in the early stages of ice encasement.     

Variable increases in cold hardiness induced in winter rape by plant growth regulators

Triazole and conventional growth regulators were tested for their ability to extend cold hardiness and improve the winter survival of winter rape (Brassica napus L.). Winter rape plants were grown in the field (Ottawa 45°23 N) and in growth cabinets. Plant growth regulators (PGRs) were applied during the early vegetative stage and the plants were allowed to cold harden. Cold-hardened plants from the field and cabinet were subjected to freezing and ice encasement tests in the laboratory. Some of the triazole PGRs reduced plant size by limiting cell expansion and increasing cell numbers. While cold hardiness and ice encasement tolerance were increased by some growth regulators, these effects were not consistent with time nor were they reflected in increased winter survival. Natural cold hardening may have eclipsed the PGR-induced hardening.PGR contribution number 1381.     

Effects of salicylic acid and cold on freezing tolerance in winter wheat leaves

The effects of salicylic acid (SA) (0.01, 0.1 and 1 mM) and cold on freezing tolerance (freezing injury and ice nucleation activity) were investigated in winter wheat (Triticum aestivum cv. Dogu-88) grown under control (20/18 °C for 15, 30 and 45-day) and cold (15/10 °C for 15-day, 10/5 °C for 30-day and 5/3 °C for 45-day) conditions. Cold acclimatisation caused a decrease of injury to leaf segments removed from the plants and subjected to freezing conditions. Exogenous SA also decreased freezing injury in the leaves grown under cold (15/10 °C) and control (15 and 30-day) conditions. Cold conditions (10/5 and 5/3 °C) caused an increase in ice nucleation activity by apoplastic proteins, which were isolated from the leaves. For the first time, it was shown that exogenous SA caused an increase in ice nucleation activity under cold (15/10 and 10/5 °C) and control conditions. These results show that salicylic acid can increase freezing tolerance in winter wheat leaves by affecting apoplastic proteins.     

WEIGHTS OF SOME PALAEARCTIC MIGRANTS IN SOUTHERN UGANDA

Between March 1966 and May 1968 Palaearctic passerines were mist-netted in thick bush and lightly wooded savannah habitats near Kampala, on the northern shore of Lake Victoria. This paper reports weights of the seven principal species involved. Most migrants appeared to be in a lean condition during the winter months, when weights were relatively low and varied little in each species. Birds were not particularly light on arrival. In fact, autumn Garden Warblers Sylvia borin and Willow Warblers Phylloscopus trochilus were sometimes markedly heavy, and for the former species there was some evidence that the individuals concerned were passage migrants. Autumn weights of Swallows Hirundo rustica, Reed Warblers Acrocephalus scirpaceus and Yellow Wagtails Motacilla flava were similar to those recorded in winter. The mean weight of all species rose during late March or early April. Although most Garden Warblers and Willow Warblers trapped at the time of spring migration were within the normal winter weight range, many Acrocephalus warblers and the majority of Sand Martins Riparia riparia and Yellow Wagtails were rather heavy. Spring weights 40% or more above mean winter weight were not uncommon in the Sedge Warbler Acrocephalus schoenobaenus, but were recorded only occasionally in other species. Although most passerine migrants evidently left Kampala with substantial fat reserves, it was concluded that a considerable number of warblers departed at rather low weight. High spring weights were mainly confined to a period of two or three weeks in each of the warbler species. Locally wintering Acrocephalus warblers must have attained full premigratory weights within three weeks, and a number of spring retraps showed substantial gains at minimum mean rates of between 0–1 and 0–35 g per day. Most heavy Garden Warblers were probably on passage. Significant correlations between weight and wing-length were obtained for all species investigated, regressions of weight on wing-length being in the range 011-0-25 g/mm. Spring weights are briefly compared with data from Nigeria, and the northward migration of passerines from Lake Victoria is discussed.     

Potential for bowhead whale entanglement in cod and crab pot gear in the Bering Sea

Bowhead whales (Balaena mysticetus) of the western Arctic stock winter in ice‐covered continental shelf regions of the Bering Sea, where pot fisheries for crabs (Paralithodes and Chionoecetes spp.) and Pacific cod (Gadus macrocephalus) pose a risk of entanglement. In the winter of 2008–2009 and 2009–2010 the spatial distribution of 21 satellite tagged bowhead whales partially overlapped areas in which pot fisheries for cod and blue king crab (Paralithodes platypus) occurred. However, these fisheries ended before whales entered the fishing areas, thus avoiding temporal overlap. A fishery for snow crab (Chionoecetes opilio) typically runs from January to May and provides the greatest potential for bowhead whales to encounter active pot gear. Tagged whales did not enter the area of the snow crab fishery during this study and generally remained in areas with >90% sea ice concentration, which is too concentrated for crab boats to penetrate. Pack ice sometimes overruns active fishing areas, resulting in lost gear, which is the most likely source of entanglement. The western Arctic stock of bowhead whales was increasing as of 2004; as such, incidental mortality from commercial pot fisheries is probably negligible at this time. Regardless, entanglement may increase over time and should be monitored.     

THE PROCESSING OF UNCONFINED,NATURALLY ENTRAINED LEAVES IN TWO LOW-ORDER SOUTHERN AFRICAN MOUNTAIN STREAMS

Summary

Breakdown of unconfined and confined leaves of the riparian tree Brabejum stellatifolium L. was investigated in two low order mountain streams (Window Stream and Langrivier) in the southwestern Cape, South Africa. At both sites, 5000 unconfined leaves were released in April and another 5000 in December 1990. The leaves were marked and half of them presoaked in river water prior to release. Leaves reaching a net 100 m below the point of release were removed periodically. On termination of the experiments, the study reaches were cleared of the remaining marked leaves. The distance which these leaves had travelled, and the retentive feature at which they were found were recorded. All leaves were weighed after recovery. In December 1990, leaves confined in coarse-mesh bags were placed on the stream beds of the two sites. Decay rates of unconfined leaves differed between streams and seasons but not between wetted and dry leaves or those of contrasting initial size and weight. In winter, breakdown of unconfined leaves was rapid (t₅₀ of 6.6d and 23.9d for Langrivier and Window Stream respectively). In summer, breakdown of unconfined leaves was slower (t₅₀ of 58d, Langrivier), but more rapid than leaves confined in mesh bags (t₅₀ of 77d, Langrivier). Distance travelled downstream had no significant effect on leaf breakdown. Different retentive features resulted in different rates of decay. Leaf weight loss in winter and summer at both sites was greatest in riffles (17–80%) and runs (21–78%). In all cases, stranded (exposed and out of the water) leaves lost the least weight (4–38%).     

COMPETITIVE SUBORDINATION OF A PIEDMONT OLD FIELD SUCCESSIONAL DOMINANT BY AN INTRODUCED SPECIES

Heterotheca latifolia Buckley has been replacing Erigeron canadensis L. as the first year dominant in Carolina-Georgia piedmont old field succession. This species of Heterotheca was introduced to this area in the 1940's. Both the Heterotheca and Erigeron species are winter annuals. We used a replacement series experiment to detect competition (interference) under a variety of greenhouse environments to seek a possible explanation for the observed changes in old field succession. Interspecific competition between Heterotheca and Erigeron plants resulted in significant reductions in the dry weight of Erigeron plants. Erigeron wilted before Heterotheca in 98.2% of observations of first-wilting in mixed-species cultures. At flowering, Erigeron plants in mixed-species cultures were shorter than Heterotheca plants. Poor seed dispersal in mixed-species communities in the field could limit Erigeron's ability to populate new areas. Greenhouse flats densely sown with equal numbers of seeds of both species resulted in a winter rosette cover of Heterotheca only. Heterotheca's assumption of the role of first year dominant in old field succession may be due to interspecific competition with Erigeron at all stages in its life cycle. These competitive effects can be observed under a variety of environmental conditions, especially those in which drought occurs.