It has been hypothesized that yellow-cedar [Chamaecyparis nootkatensis (D. Don) Spachl decline may result from root freezing injury following climate change-induced reductions in protective snow cover. To test this hypothesis, we measured the freezing tolerance and injury expression of yellow-cedar seedlings in three treatments that differed in the insulative protection they provided to soils during winter and spring: (1) full exposure to ambient temperatures (exposed treatment), (2) continuous protection from ambient temperatures via addition of perlite over pots (full protection), and (3) perlite protection only during winter and exposure to ambient temperatures during spring (partial protection). Foliage from all treatments was cold tolerant enough to prevent foliar freezing injury throughout the study period. However, on all sample dates, roots of seedlings from all treatments were only tolerant to about -5 "C -a level considerably warmer than the reported maximum cold tolerance for the species and well above the soil temperature recorded in the exposed treatment. As a result of this limited root cold tolerance, visibly uninjured roots of seedlings from the exposed treatment had significantly higher relative electrolyte leakage (REL) throughout the winter and early spring than seedlings in soil protection treatments. Seedlings from the exposed treatment also had significantly higher foliar REL values and greater visual foliar injury than seedlings from the other treatments starting in early spring. For both roots and foliage, REL measurements consistently detected tissue damage before visual injury was evident. Patterns of injury from both REL and visual injury assessments showed the same pattern: damage began with freezing injury to roots and subsequently became evident as foliar browning after spring temperatures increased. All seedlings in the exposed treatment eventually had 100% fine root damage and died. This progression of initial root damage followed by foliar browning and mortality after the onset of warming conditions is consistent with reports of yellow-cedar decline symptom development in the field.
Numerous anthropogenic factors can deplete calcium (Ca) from forest ecosystems. Because an adequate supply of Ca is needed to support fundamental biological functions, including cell membrane stability and stress response, the potential for Ca deficiency following the individual, cumulative, or potentially synergistic, influences of anthropogenic factors raises important questions concerning organism and ecosystem health. Past work has shown that one Ca-depleting factor (foliar acid mist exposure) reduces concentrations of biologically important membrane-associated Ca (mCa) from red spruce foliar cells, destabilizes these cells, and results in their increased susceptibility to the freezing injury responsible for red spruce decline in northeastern U.S. montane eco-systems. Data presented here indicate that these same disruptions can occur for other tree species and that soilbased Ca manipulation can also alter critical mCa pools. Considering the unique role Ca plays in the physiological response of cells to environmental change and stress, we hypothesize that depletion of biologically available Ca (e.g., mCa) could result in a scenario similar to recognized immune deficiency syndromes in animals. A hypothetical pathway through which anthropogenically induced Ca deficiencies could predispose plants, and possibly animals, to exaggerated injury following exposure to environmental stress is presented, and the potential implications of this scenario to ecosystem health are discussed.
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·ha1·year1 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.
In fall (November 2005) and winter (February 2006), we collected current-year foliage of native red spruce (Picea rubens Sarg.) growing in a reference watershed and in a watershed treated in 1999 with wollastonite (CaSiO(3), a slow-release calcium source) to simulate preindustrial soil calcium concentrations (Ca-addition watershed) at the Hubbard Brook Experimental Forest (Thornton, NH). We analyzed nutrition, soluble sugar concentrations, ascorbate peroxidase (APX) activity and cold tolerance, to evaluate the basis of recent (2003) differences between watersheds in red spruce foliar winter injury. Foliar Ca and total sugar concentrations were significantly higher in trees in the Ca-addition watershed than in trees in the reference watershed during both fall (P=0.037 and 0.035, respectively) and winter (P=0.055 and 0.036, respectively). The Ca-addition treatment significantly increased foliar fructose and glucose concentrations in November (P=0.013 and 0.007, respectively) and foliar sucrose concentrations in winter (P=0.040). Foliar APX activity was similar in trees in both watersheds during fall (P=0.28), but higher in trees in the Ca-addition watershed during winter (P=0.063). Cold tolerance of foliage was significantly greater in trees in the Ca-addition watershed than in trees in the reference watershed (P<0.001). Our results suggest that low foliar sugar concentrations and APX activity, and reduced cold tolerance in trees in the reference watershed contributed to their high vulnerability to winter injury in 2003. Because the reference watershed reflects forest conditions in the region, the consequences of impaired physiological function caused by soil Ca depletion may have widespread implications for forest health.
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