Influence of insulating dead leaves and low temperatures on water balance in an Andean giant rosette plant

Abstract. The influence of insulating dead leaves on water balance in Espeletia timotensis Cuatr., an Andean caulescent giant rosette plant, was studied under field and laboratory conditions. Removal of the dead leaf layer surrounding the stem changed the pattern of diurnal stem temperature variation and produced transient and permanent effects on water balance. The pattern of liquid water flow resistance increase at low stem temperatures suggested that much of the water flow in the stem was through living membranes, probably those of the pith cells. The pith was determined to be an important source of stored water for daily transpirational needs. The lethal effects of dead leaf removal were attributed to one or more of the following causes: (1) inhibition of pith recharge by subfreezing stem temperatures; (2) embolisms in stem xylem; (3) freezing injury to pith tissue. The results suggested that an insulating layer of marcescent leaves and the presence of an internal water reservoir closer to the rosette than the soil water are important adaptations for maintenance of a favourable water balance in tropical alpine habitats where freezing temperatures occur regularly but last only a few hours.     

The role of capacitance in the water balance of Andean giant rosette species

Abstract Pith water storage capacity and its role in plant-water relations were studied in seven giant rosette species of the genus Espeletia from the Venezuelan Andes. Readily available water from the pith was calculated to be capable of sustaining mean transpiration for up to 2.5 h. The relative importance of water stored in the pith, however, differed among species. The species that grow in the higher and colder environments tended to have a greater capacitance than the species that grow in the lower and less extreme environments. The pith volume per unit leaf area (PV/LA) was found to be a good indicator of the relative water storage capacity of the adult individuals of each species. Diurnal fluctuations in leaf water potential were not as pronounced in the species with higher PV/LA values. The species-specific PV/LA was highly correlated with the leaf turgor loss point and with the total resistance to water flow from soil to leaves. These results suggested that species-specific capacitance in the genus Espeletia is a response to temperature-limited soil water availability and that cold tropical environments with frequent subfreezing temperatures tend to select for high water storage capacity in giant rosette plants.     

Freezing tolerance and avoidance in high-elevation Hawaiian plants

Freezing resistance mechanisms were studied in five endemic Hawaiian species growing at high elevations on Haleakala volcano, Hawaii, where nocturnal subzero (°C) air temperatures frequently occur. Extracellular freezing occurred at around -5°C in leaves of Argyroxiphium sandwicense and Sophora chrysophylla, but these leaves can tolerate extracellular ice accumulation to -15°C and -12°C, respectively. Mucilage, which apparently acted as an ice nucleator, comprised 9 to 11% of the dry weight of leaf tissue in these two species. Leaves of Vaccinium reticulatum and Styphelia tameiameiae were also found to tolerate substantial extracellular freezing. Dubautia menziesii, on the other hand, exhibited the characteristics of permanent supercooling; a very rapid decline in liquid water content associated with simultaneous intracellular and extracellular freezing. However, in those species that tolerate extracellular freezing, the decline in liquid water content during freezing is relatively slow. Osmotic potential was lower at pre-dawn than at midday in four of the species studied. Nocturnal production of osmotically active solutes may have helped to prevent intracellular freeze dehydration as well as to provide non-colligative protection of cell membranes. Styphelia tameiameiae supercooled to -9·3°C and tolerated tissue freezing to below -15°C, a unique combination of physiological characteristics related to freezing. Tolerance of extracellular ice formation after considerable supercooling may have resulted from low tissue water content and a high degree of intracellular water binding in this species, as determined by nuclear magnetic resonance studies. The climate at high elevations in Hawaii is relatively unpredictable in terms of the duration of subzero temperatures and the lowest subzero temperature reached during the night. It appears that plants growing in this tropical alpine habitat have been under selective pressures for the evolution of freezing tolerance mechanisms.     

Growth rates, reproductive phenology, and pollination ecology of Espeletia grandiflora (Asteraceae), a giant Andean caulescent rosette

From March 2001 to December 2002, we studied the reproductive phenology, pollination ecology, and growth rates of Espeletia grandiflora Humb. and Bonpl. (Asteraceae), a giant caulescent rosette from the Páramos of the Eastern Andes of Colombia. Espeletia grandiflora was found to be predominantly allogamous and strongly self-incompatible. Bumblebees (Bombus rubicundus and B. funebris) were the major pollinators of E. grandiflora, although moths, hummingbirds, flies, and beetles also visited flowers. Inflorescence development began in March and continued through August to September. Plants flowered for 30 - 96 days with a peak from the beginning of October through November. The percentage of flowering plants strongly differed among size classes and between both years. Seed dispersal occurred as early as September through May of the following year. The average absolute growth rate for juveniles and adults rate was 7.6 cm/year. Given the scarcity of floral visitors at high altitudes due to climatic conditions, we suggest that even small contributions from a wide range of pollinators might be advantageous for pollination of E. grandiflora. Long-term studies on different populations of E. grandiflora are required to determine if the high growth rates are representative, to quantify the variation in the flowering behavior within and among populations, and to establish if nocturnal pollination is a trait that is exclusive to our population of E. grandiflora.     

Frost avoidance and freezing tolerance in Afroalpine 'giant rosette' plants

Abstract. The large leaf rosettes of the so-called 'giant rosette' plants which inhabit the alpine region of tropical mountains are composed of a great number of adult leaves surrounding a central cone of developing leaves. Upon onset of the nocturnal frost period the adult leaves nyctinastically bend inwards and form a night-bud around the central leaf bud. The insulating effect of the night-bud was analysed in four species: Senecio keniodendron, Senecio brassica. Lobelia keniensis and Lobelia telekii which grow on Mt Kenya (Kenya). Freezing is avoided by a delay of cooling which is sufficient until rewarming by the next day's sunshine. A consequence of this delay is that the temperature in the nocturnal bud remains higher than that of the outer leaves which are often stiffly frozen after cold nights. Only one freezing point was detected on the leaf temperature recordings. Depending on the water state, the freezing points were in the range — 1°C to —4.2°C. Sucrose, amounting to 38% of the leaf dry weight, may act as a cryoprotectant for the cell membranes. Frost hardiness of the leaves, as determined with a laboratory method, was sufficient to enable the plants to survive during the nocturnal frost temperatures as measured in the field.     

Freezing avoidance by deep undercooling of tissue water in winter-hardy plants

Freezing avoidance by deep undercooling of tissue water to near its homogeneous nucleation temperature (approximately −40 °C) has recently been shown to be an important survival mechanism in reproductive and vegetative parts of many winter-hardy plants. Biophysical experiments which support the concept of undercooling of the tissue water include thermal analyses, nuclear magnetic resonance spectroscopy, and low-temperature microscopy of tissue freezing. All these experiments suggest that in plant parts that undercool, tissue water is compartmentalized and is not removed to extracellular ice as tissue temperature declines. When freezing takes place at low temperature, it occurs rapidly and appears to be intracellular, resulting in instant death. Analyses of freezing and injury of winter-hardy plants at the northern limits of the deciduous forest in North America and near timberline in the Rocky Mountains of the western United States indicate that freezing survival by deep undercooling is an important factor in limiting plant distribution.     

Geographic variation in winter freezing susceptibility in the eggs of the European pine sawfly (Neodiprion sertifer)

Abstract 1 The European pine sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae), frequently defoliates Scots pine (Pinus sylvestris L.) forests in northern Europe. It overwinters as an egg. It has been proposed that the high egg mortality caused by low winter temperatures limits the occurrence of outbreaks to the southern part of Fennoscandia. 2 In this study, variation in freezing avoidance by egg supercooling between four Finnish populations (originating between latitudes 60°N and 69°N) of N. sertifer was tested by differential thermal analysis. Offspring of 20 females within each population were selected for the study. The freezing avoidance of parasitized eggs was also examined. 3 The northernmost Inari population was found to be the cold hardiest, and the southernmost (Hanko) was the least hardy population. The within‐population variation between females was greatest in the population from Inari, and the next greatest in the one from Hanko. The inland populations in Eastern Finland had the smallest within‐population variation in freezing avoidance. 4 The high variation in freezing avoidance of eggs will enable N. sertifer to adapt to the predicted climate change and to spread its distribution northwards. This may also change the risk for outbreaks in this area. Parasitized eggs froze at higher temperature than healthy eggs. This observation indicates that N. sertifer may experience reduced egg parasitism in certain winter climate conditions.     

Some characteristics of freezing avoidance in two osmerids, rainbow smelt and capelin

Winter-acclimatised rainbow smelt Osmerus mordax have elevated levels of glycerol and trimethylamine oxide (TMAO), and a protein antifreeze, all of which are lost in summer. No seasonal differences were found in glycerol-3-P-dehydrogenase, glycerol-3-phosphatase and glutamate dehydrogenase, which are involved in the synthesis of glycerol or its precursors. Although glycerol production requires NADH, no large seasonal differences were found in the activities of glucose-6-P-dehydrogenase and 6-phosphogluconate dehydrogenase (pentose phosphate pathway), malate dehydrogenase and aspartate aminotransferase (malate-aspartate shuttle) or in the NAD : NADH ratio. Thus, other regulatory mechanisms must be involved. Capelin Mallotus villosus collected from mid water in the Barents Sea in September (water temperatures +0·4 to −1·6°C) had only negligible amounts of glycerol, and no protein antifreeze, although serum TMAO levels (44 mm) were unusually high for a teleost. Capelin appear to avoid freezing largely by avoiding water layers with temperatures below their body freezing point, and by occasional supercooling.     

Freezing behaviours in wintering Cornus florida flower bud tissues revisited using MRI

How plant tissues control their water behaviours (phase and movement) under subfreezing temperatures through adaptative strategies (freezing behaviours) is important for their survival. However, the fine details of freezing behaviours in complex organs and their regulation mechanisms are poorly understood, and non‐invasive visualization/analysis is required. The localization/density of unfrozen water in wintering Cornus florida flower buds at subfreezing temperatures was visualized with high‐resolution magnetic resonance imaging (MRI). This allowed tissue‐specific freezing behaviours to be determined. MRI images revealed that individual anthers and ovules remained stably supercooled to ?14 to ?21 °C or lower. The signal from other floral tissues decreased during cooling to ?7 °C, which likely indicates their extracellular freezing. Microscopic observation and differential thermal analyses revealed that the abrupt breakdown of supercooled individual ovules and anthers resulted in their all‐or‐nothing type of injuries. The distribution of ice nucleation activity in flower buds determined using a test tube‐based assay corroborated which tissues primarily froze. MRI is a powerful tool for non‐invasively visualizing unfrozen tissues. Freezing events and/or dehydration events can be located by digital comparison of MRI images acquired at different temperatures. Only anthers and ovules preferentially remaining unfrozen are a novel freezing behaviour in flower buds. Physicochemical and biological mechanisms/implications are discussed.     

<Emphasis Type="Italic">Espeletia</Emphasis> giant rosette plants are reliable biological indicators of time since fire in Andean grasslands

Páramo grasslands in the tropical Andes are fire-prone ecosystems and an understanding of their fire ecology is fundamental to biodiversity conservation and ecosystem management. Fire registers are normally impractical in these remote, cloud-covered landscapes, but Espeletia giant rosette plants have been proposed as biological indicators of time since fire in páramos. Espeletia giant stem rosettes tolerate fire well, protecting apical buds in at the heart of their leaf rosettes, and for some species, germination is known to be enhanced by fire. As the plant grows, its dead leaves remain attached to the stem, but fire removes these and resets the “leaf clock”. This study uses a unique register of fires in one Ecuadorian páramo to assess the robustness of this biological indicator. Dead leaf cover on Espeletia pycnophylla giant rosette plants was measured in fifteen different sites with known fire dates from 2000 to 2014. The growth rates of plants at four different elevations were measured over a 2-year period and used to estimate time since fire based on dead leaf cover in the known sites. Estimates were accurate to ± 2 year. Thus, where fire records are missing, relatively easy measurements of growth rates and dead leaf cover of Espeletia giant rosette plants can provide reliable estimates across a wide range of times since fire. This approach has value for direct investigations into fire ecology but also for studies in which controlling for fire dynamics is necessary to reveal underlying patterns. Therefore, this approach also offers a means to obtain better information on other landscape-scale processes such as the impact of climate change on biodiversity or the provision of ecosystem services.     

Freezing avoidance by supercooling in Olea europaea cultivars: the role of apoplastic water,solute content and cell wall rigidity

Plants can avoid freezing damage by preventing extracellular ice formation below the equilibrium freezing temperature (supercooling). We used Olea europaea cultivars to assess which traits contribute to avoid ice nucleation at sub‐zero temperatures. Seasonal leaf water relations, non‐structural carbohydrates, nitrogen and tissue damage and ice nucleation temperatures in different plant parts were determined in five cultivars growing in the Patagonian cold desert. Ice seeding in roots occurred at higher temperatures than in stems and leaves. Leaves of cold acclimated cultivars supercooled down to ?13 °C, substantially lower than the minimum air temperatures observed in the study site. During winter, leaf ice nucleation and leaf freezing damage (LT₅₀) occurred at similar temperatures, typical of plant tissues that supercool. Higher leaf density and cell wall rigidity were observed during winter, consistent with a substantial acclimation to sub‐zero temperatures. Larger supercooling capacity and lower LT₅₀ were observed in cold‐acclimated cultivars with higher osmotically active solute content, higher tissue elastic adjustments and lower apoplastic water. Irreversible leaf damage was only observed in laboratory experiments at very low temperatures, but not in the field. A comparative analysis of closely related plants avoids phylogenetic independence bias in a comparative study of adaptations to survive low temperatures.     

Cold resistance mechanisms in high desert Andean plants

Freezing tolerance and freezing avoidance were studied, during the growing season, in plant species from two different elevations (3200 m and 3700 m) in a desert region of the high Andes (29° 45S, 69° 59W) in order to determine whether there was a relationship between plant height and cold resistance mechanisms. Freezing injury and supercooling capacity were determined in plants of different height, from ground-level (<20 cm tall) to tall shrubs (27–90 cm). All ground-level plants showed freezing tolerance as the main mechanism for resistance to freezing temperatures. Tall shrubs avoided freezing temperatures, mainly through supercooling. Supercooling was only present in plants occupying the lower elevation (i.e., 3200 m). Both avoidance and tolerance mechanisms are present in a single genus (i.e., Adesmia).     

甜杨6-磷酸葡萄糖脱氢酶在抗冻性低温诱导中的作用

对-20℃低温锻炼及脱锻炼过程中甜杨(Populus suaveolens)幼苗的G6PDH、SOD和POD活性、MDA含量和半致死温度(LT50)进行了测定和分析.结果发现,低温锻炼在一定程度上提高了幼苗6-磷酸葡萄糖脱氢酶(G6PDH)、SOD和POD活性,降低了MDA含量和幼苗半致死温度(LT50).另外,将幼苗放回常温(脱锻炼)2 d能引起幼苗的G6PDH、SOD和POD活性的显著下降,并使LT50和MDA含量的迅速回升.结果表明,低温锻炼中G6PDH活性的增加有助于SOD和POD活性的提高,进而对幼苗的LT50和MDA含量的降低有明显的促进作用,G6PDH可能参与了SOD和POD活性的调节和抗冻性的低温诱导.     

Phenotype matching and inbreeding avoidance in African elephants

Perhaps the most important 'decision' made by any animal (or plant) is whether to disperse--leave kith and kin, or remain with the familiar and related. The benefits of staying at home are obvious, so dispersal requires an explanation--and the most popular is that dispersal functions to avoid inbreeding depression. Strong support comes from the observation that dispersal is so often sex biased. Simply put, all else being equal members of both sexes should prefer to remain philopatric, but this would lead to inbreeding depression so members of one sex have to disperse. In principle, this link between inbreeding depression and sex-biased dispersal could be broken if individuals recognize close kin and avoid mating with them. Archie et al. (2007) provide one of the most compelling analyses to date of the interaction among inbreeding avoidance, kin recognition and mating strategies in any mammal, clearly showing that elephants recognize even close paternal kin and avoid mating with them. Their important results illuminate the subtleties of elephant inbreeding avoidance as well as illustrate the difficulty of arriving at definitive answers to questions about the evolution of dispersal behaviour.     

使用冷冻方法防治昆虫标本虫害

皮蠹幼虫对昆虫标本的蛀蚀是我国北方地区标本保藏时需要注意的首要问题。经12次的试验观察表明,花斑皮蠹幼虫TrogodermavariabileBallion在冰柜中放置位置不同其冷冻致死率亦不相同置于冰柜表层的,死亡率介于0～50%;上层的死亡率为95%～100%;中上层及中层死亡率达100%。放置在表层及上层的皮蠹幼虫,在经2d以上的冷冻处理后部分个体出现复活。因此,对那些原先放置上层及表层的标本,第1次冷冻结束后,间隔数天应再进行第2次冷冻,以提高和巩固冷冻杀虫效果。经3年的实践证明,采用冷冻方法治理皮蠹幼虫为害效果明显,可以推广普及。     

Cold adaptation in insects of Central Yakutia

The mode of cold hardening was for the first time assessed for 20 insect species living in the extremely cold climate of Yakutia. All insects tested were found to adapt through freeze tolerance, producing ice-nucleating agents that cause the hemolymph to freeze at high subzero temperatures. For the first time ice-nucleating agents were demonstrated in Lepidoptera. Pieris rapae exemplified the possibility of switchover in the survival strategy depending on the climatic conditions.     

Predator avoidance in mummichog larvae from a polluted habitat

Previous work has shown that adult mummichogs ( Fundulus heteroclitus ) from a polluted site (Piles Creek, PC, New Jersey) were more vulnerable to predation by blue crabs than fish from a more pristine site (Tuckerton, TK, New Jersey). The present study was conducted to compare the spontaneous activity, swimming performance and predator avoidance of fish of the two populations at early life stages. When raised in synthetic salt water, both newly hatched and 1–month-old mummichog larvae from TK had greater spontaneous activity, swimming performance, and stamina than those from PC. In comparison to TK, PC larvae were less vulnerable to predation by yearling mummichogs at 1 week old, but were more vulnerable at 1 month old. Variation in behaviour and swimming ability of TK and PC larvae accounted for the differences in vulnerability to predation. TK larvae employed greater swimming and irregular movement, which was more successful for survival of older larger larvae. In contrast, PC larvae hid, often remaining motionless, which was advantageous for survival of young smaller larvae. Contaminants had been assumed responsible for the behavioural differences in the PC population. However, since differences were seen in larvae raised in synthetic salt water, it is possible that maternally transferred neurotoxicants (possibly delayed effects) and/or inherent factors also play a role.     

The role of chlorpheniramine in inhibitory avoidance in Danio vevio

The action of chlorpheniramine in learning and memory in Danio rerio suggested that 1 mg kg⁻¹ produced an amnesiac effect in inhibitory avoidance in this species.