No abstract

To provide baseline data for physiological studies of extreme low-temperature (LT) tolerance in boreal conifers, we profiled LT stress responses, liquid nitrogen (LN(2))-quench tolerance, and sugar concentrations in foliage of boreal-temperate species pairs in the genera Abies, Picea and Pinus, growing in an arboretum in a temperate oceanic climate from August 2006 through April 2007. The boreal species acclimated more rapidly and deeply than the temperate species, acquiring LN(2)-quench tolerance by late November, despite unusually warm conditions throughout the autumn and early winter. Maximum LT tolerance in the temperate species was in the -25 to -35 degrees C range, and was reached only after a period of freezing temperatures in late January and February. During LT acclimation in the temperate species, sigmoid temperature-relative electrolyte leakage (REL) curves shifted toward lower temperatures, whereas in boreal species there was both a temperature shift and a lowering of the maximum REL until it fell below a threshold associated with irreversible injury. These differences may reflect differences in mechanisms of LT acclimation and LT tolerance. The concentrations of total and individual sugars did not show a clear pattern that could differentiate the boreal and temperate groups. Raffinose and, in three of the six species, stachyose showed the closest association with LT tolerance. Sugar concentrations, principally sucrose, decreased during mild weather, perhaps because of respiratory losses or phloem export, and increased after periods of freezing temperatures. Low-temperature acclimation in boreal species appears to follow a rigid program that may affect their ability to avoid excessive respiratory losses in the event of continued climate warming in boreal regions.

Because they maintain green foliage throughout the winter season, evergreen conifers may face special physiological challenges in a warming world. We assessed the midwinter low-temperature (LT) tolerance of foliage from eight temperate and boreal species in each of the genera Abies, Picea, and Pinus growing in an arboretum in Trondheim, Norway, using relative electrolyte leakage (REL) as an index of cell injury. Relatively LT sensitive species came from temperate coastal and Mediterranean environments and displayed a well-defined sigmoidal response to LT stress, with LT 50 ranging from -27 to -38°C. Species originating from boreal regions were not lethally stressed by slow freezing to temperatures as low as -80°C, while species from temperate mountains and continental interiors displayed intermediate responses, with LT50s ranging from -33 to -44°C. Further evaluation of one sensitive and one insensitive species in each genus showed that boreal species can survive quenching in liquid nitrogen at -196°C provided they are first slowly cooled to -30°C or lower. Quantitative image analysis of color changes resulting from LT stress followed by exposure to light showed that foliage from nonlethally stressed boreal species developed mild to moderate chlorosis while more sensitive species developed a mixture of chlorosis and necrosis, with significant necrosis occurring mainly at temperatures resulting in REL of 50% or more. Sensitive and insensitive trees differed significantly in total raffinose, sucrose, and total sugar concentrations, and raffinose and sucrose correlated significantly with LT 50 within the sensitive group.

We evaluated the influence of protracted low-level nitrogen (N) fertilization on foliar membrane-associated calcium (mCa), sugar and starch concentrations, membrane stability, winter cold tolerance, and freezing injury of red spruce (Picea rubens Sarg.) trees growing in six experimental plots on Mount Ascutney, Vermont. For 12 consecutive years before this evaluation, each plot received one of three treatments: 0, 15.7, or 31.4 kg N·ha–1·year–1 supplied as NH4Cl. In comparison with trees from control plots, the current-year foliage of trees from N-addition plots had lower mCa concentrations, higher levels of electrolyte leakage, reduced cold tolerance, and greater freezing injury. Levels of mCa, membrane stability, and cold tolerance did not differ between N treatments, but trees in high-N treated plots experienced greater freezing injury. Although no differences in carbohydrate nutrition were detected in September, foliar sugar and starch concentrations from trees in N-treated plots were higher than control plot trees in January. We propose that foliar mCa deficiencies reduced cell membrane stability, decreased cold tolerance, and increased freezing injury for trees in N addition plots relative to controls. Declines in mCa may also help account for increases in respiration previously measured. Because soil, root, and mycorryhizal conditions were not evaluated, it is unknown how treatment-induced changes in these compartments may have influenced the alterations in foliar mCa and physiological parameters measured in this study.

We documented 3 to 14°C of dehardening in current-year foliage of 10 mature, montane red spruce (Picea rubens Sarg.) trees during a natural thaw from 12 to 21 January 1995. Mean cold tolerance was about -47°C before the onset of thaw conditions, and individuals ranged from -38 to -52OC. After 3 days of thaw, mean cold tolerance dropped to -39"C, with a range of -32 to -44°C. Trees did not regain prethaw levels of cold tolerance until sometime between 31 January and 9 February, or 10 to 20 days after subfreezing temperatures resumed. The least cold tolerant tree was at risk of injury when temperature at the field site fell to an estimated -33.8"C on 6 February, and this same tree developed noticeably more injury than other trees when injury symptoms developed in late March. No evidence of dehardening was found in balsam fir (Abies balsamea (L.) Mill.) trees from the same stand. All red spruce trees also showed the potential for net assimilation of carbon during the thaw, as determined by measurement of photosynthetic capacity under laboratory conditions. From the abrupt and substantial dehardening and persistence of the dehardened state, we conclude that dehardening during periods of warm weather may be a significant factor in freezing injury and decline of montane red spruce populations.