Emiko Maruta

Effects of high light and low temperature during harsh winter on needle photodamage of Abies mariesii growing at the forest limit on Mt. Norikura in Central Japan

Abstract Coniferous evergreen firs Abies mariesii (A. mariesii) growing at the forest limit (near 2500 m altitude) on Mt. Norikura (36°61′N, 137°33′E, 3026 m altitude) in Central Japan are exposed to severe winter-stresses. Under such conditions, only the abaxial side of A. mariesii needles changes from green to reddish–brown in early spring, resulting in the death of the needles. Since this needle damage is only observed in shoots that protrude from the snow surface and not in those buried in snow or located at lower elevations, this phenomenon seems to be caused by the interaction of strong sunlight reflected from the snow surface and low temperature. We found that the damaged needles have increased in the de-epoxidation state of xanthophyll cycle because they contained large amounts of zeaxanthin, which appears when the leaves encounter a strong light stress, leading to the long-term down-regulation of PS II photochemistry. These results indicate that the needles acclimatize against the strong light during severe winter. Furthermore, ascorbate peroxidase (APX, E.C. 1.11.1.11) activity was found to decrease remarkably during critical subzero periods, while superoxide dismutase (SOD, E.C. 1.15.1.1) activity remains constant throughout the year. Based on these results, we discuss herein how A. mariesii growing at the forest limit of temperate zone responds to high light and low temperature in harsh winter conditions.

Patch establishment and development of a clonal plant, Polygonum cuspidatum, on Mount Fuji

Microsatellite analysis was used to investigate the patch establishment and development of Polygonum cuspidatum Sieb. et Zucc, a clonal herbaceous plant that dominates the primary succession on the southeast slope of Mount Fuji. Genotypes of P. cuspidatum in 155 patches at the study site differed from each other. This indicates that P. cuspidatum patches are initially established by seed dispersed on the bare scoria field, and not by clonal rhizome extension. Genetic differentiation was estimated using the FST values between subpopulations at the study site. There was almost no genetic differentiation between subpopulations, indicating the presence of massive gene flow. The pollen fathers of seeds and maternal genets of current‐year seedlings were inferred from the microsatellite allele composition by a simple exclusion method. The wide, random distribution of pollen fathers suggests that pollen dispersal occurs over a broad area. Maternal analysis showed a tendency for seed dispersal to be biased to the area nearby and down slope from the mother plants. Patch establishment under massive gene flow may result from such pollen and seed dispersal. To understand the process of patch development, aerial photographs taken from 1962 to 1999 were compared, and then genets in each of 36 patches were identified from the microsatellite genotypes of P. cuspidatum shoots. The comparison of aerial photographs showed that most of the patches enlarged each year and that some neighbouring patches combined during growth. Genet analysis demonstrated a high correlation between patch area and the area of the largest genet within it, and that new genets were recruited at the patch periphery. These findings indicate that both vegetative and sexual reproduction, i.e. rhizome extension and the establishment of new seedlings, contribute to the development of P. cuspidatum patches.

Diurnal changes in needle gas exchange in alpine Pinus pumila during snow‐melting and summer seasons

Pinus pumila (Pallas) Regel. is a dominant dwarf tree in alpine regions of Japan. The possible factors limiting the net photosynthetic rate (Pn) of the needles of P. pumila were examined in the snow-melting (May) and the summer (August) seasons. In August, in situ maximum Pn was 20 μmol kg−1 needle s−1 in the current-year needles and 25 μmol kg−1 needle s−1 in the 1-year-old needles. Diurnal trends of Pn in August were positively related to fluctuations in photosynthetic photon flux density (PPFD) and no midday depression of Pn was found, indicating that a decrease in PPFD rather than an increase in needle-to-air vapor pressure deficit (ΔW) might cause the reduction of Pn. Both stomatal conductance (gs) and Pn were lower in May than in August. In May, Pn and gs were almost zero in the morning, but gradually increased with decreasing ΔW, reaching maximum Pn values (4 μmol kg−1 needle s−1) and gs (60 mmol kg−1 needle s−1) at 16.00 hours. The daytime Pn in May was positively related to gs. Relative water content in the exposed needles above the snow in May was 83%, which was far above the lethal level. This indicates that the water flow from stems or soils to needles was enough to compensate for a small amount of water loss due to the low gs in May, although the water supplied to needles would be impeded by the low temperatures. Thus, the reduced gs in May would be important for avoiding needle desiccation, and would result in a reduced Pn.

Desiccation-tolerance of Fagus crenata Blume Seeds from Localities of Different Snowfall Regime in Central Japan

In beech (Fagus crenata Blume) forests on the Pacific Ocean side in Central Japan, snowpack depth is little and xeric conditions may prevail in winter, in contrast to heavy snow in beech forests on the Japan Sea side. The effects of such conditions during winter on the viability of beech seeds were studied at a beech forest on the Pacific Ocean side. Thickness and weight ratio of pericarp of beech seeds were significantly greater in populations on the Pacific Ocean side compared to the Japan Sea side, this apparently being related to snowpack depth in winter. During the initial stage of seed drying, the drying rate of seeds from the Fuji population (the Pacific Ocean side) was less than that from the Sumon population (the Japan Sea side), possibly due to the thicker pericarp. Germination percentage of seeds which dried to a water content as low as 7% d.w. did not decrease for either population, indicating desiccation tolerance of beech seeds. In the beech forest (University Forest at Yamanaka) on the Pacific Ocean side, water content of seeds sown below litter from both populations remained sufficient for viability during winter. Germinating seeds from the populations on the Japan Sea side were more desiccation-sensitive in early spring, compared to those from the Fuji population, partially due possibly to differences in pericarp thickness.

Needle traits of an evergreen, coniferous shrub growing at wind‐exposed and protected sites in a mountain region: does Pinus pumila produce needles with greater mass per area under wind‐stress conditions?

Snow depth is one of the most important determinants of vegetation, especially in mountainous regions. In such regions, snow depth tends to be low at wind-exposed sites such as ridges, where stand height and productivity are limited by stressful environmental conditions during winter. Siberian dwarf pine (Pinus pumila Regel) is a dominant species in mountainous regions of Japan. We hypothesized that P. pumila produces needles with greater mass per area at wind-exposed sites than at wind-protected sites because it invests more nitrogen (N) in cell walls at the expense of N investment in the photosynthetic apparatus, resulting in increased photosynthetic N use efficiency (PNUE). Contrary to our hypothesis, plants at wind-exposed site invested less resources in needles, as exhibited by lower biomass, N, Rubisco and cell wall mass per unit area, and had higher photosynthetic capacity, higher PNUE and shorter needle life-span than plants at a wind-protected site. N partitioning was not significantly different between sites. These results suggest that P. pumila at wind-exposed sites produces needles at low cost with high productivity to compensate for a short leaf life-span, which may be imposed by wind stress when needles appear above the snow surface in winter.

Seasonal changes in the excess energy dissipation from Photosystem II antennae in overwintering evergreen broad-leaved trees Quercus myrsinaefolia and Machilus thunbergii.

We monitored chlorophyll (Chl) fluorescence, pigment concentration and the de-epoxidation state of the xanthophyll cycle (DPS(1)) in two warm temperate broad-leaved evergreen species (Quercus myrsinaefolia and Machilus thunbergii). Reduction of the maximal quantum yield of Photosystem II (PSII) (calculated from Fv/Fm, variable to maximal Chl a fluorescence) and retention of a high DPS were observed in both species in the winter, and can be interpreted as acclimation to winter. In particular, the acclimation of PSII in these species can be chiefly attributed to thermal dissipation, which is correlated with the retention of high zeaxanthin. Furthermore, we attempted to divide the fate of the absorbed light energy by the PSII antennae into three components: (i) PSII photochemistry (represented by its quantum yield, ΦPSII), (ii) dissipation by down-regulation via non-photochemical quenching (ΦNPQ) and (iii) other non-photochemical processes (ΦONP). The estimated energy allocation of the absorbed light indicated that the proportion of ΦPSII decreased, whereas that of ΦNPQ+ΦONP increased during winter. This result suggests that the excess energy absorbed in the PSII complexes is safely dissipated from the PSII antennae. Based on these results, we conclude that thermal dissipation from the PSII antennae plays an important role in two overwintering broad-leaved evergreen trees growing in Japan.

Photoprotective mechanisms against winter stresses in the needles of Abies mariesii grown at the tree line on Mt. Norikura in Central Japan

Evergreen fir Abies mariesii growing at the tree line (near 2 500 m altitude) on Mt. Norikura (36°61′N, 137°33′E, 3 026 m altitude) in Central Japan is exposed to harsh winter stresses. To protect against these stresses, the deep-oxidation state of the xanthophyll cycle pigments increased, because the needles contained large amounts of zeaxanthin, which resulted in an increase of non-radiative thermal dissipation from the antenna system. Not only the antenna system but also the inactivated photosystem (PS) 2 reaction centre (RC) might contribute to the heat dissipation of absorbed excess photon energy. In addition, a decrease in the PS2 activity during winter was derived from the degradation of the PS2 RCs. Thus the needles acclimated to the strong sunlight during the harsh winter. Under such conditions, only the abaxial side of A. mariesii needles occasionally changed colour from green to reddish-brown in early spring. Since this needle damage was only observed in shoots that protruded from the snow surface, this phenomenon might be caused by the interaction between the strong sunlight reflected from the snow surface and the long period of sub-zero temperatures. We also examined how the photoprotective functions of A. mariesii growing at the tree line of a temperate zone mitigate the interactive stresses of high photon flux density and sub-zero temperature in harsh winter.

Pinus pumila Photosynthesis is Suppressed by Water Stress in a Wind-Exposed Mountain Site

Abstract Wind exposure is known to have stressful effects on plant growth, particularly at high altitudes. We studied how environmental factors affected carbon assimilation in Pinus pumila needles at a wind-exposed site. Needle gas exchange rates were determined for detached shoots in the laboratory where the needles were free from field environmental stresses, and also determined for attached shoots in the field under in situ environment. There was no difference in gas exchange characteristics determined in the laboratory between shoots from the wind-exposed and the wind-protected sites, suggesting that wind exposure did not affect the photosynthetic potential. In the field, however, the photosynthetic rate of one-year-old needles (in situ Aarea) was significantly lower at the windexposed site than that at the wind-protected site. There was a positive correlation between the in situ Aareaand the xylem pressure potential, suggesting that water deficit caused photosynthetic suppression at the wind-exposed site. The in situ Aarea was lower at the wind-exposed site, even with the same electron transport rate and the same stomatal conductance. These results suggest that CO2 assimilation is suppressed by lower mesophyll CO2 conductance at the wind-exposed site. We conclude that the carbon gain is limited by water stress in wind-exposed regions.

Needle browning and death in the flagged crown of Abies mariesii in the timberline ecotone of the alpine region in central Japan

In the flagged crown, which is asymmetric growth formed by severe stresses during winter in alpine regions, needles of evergreen conifers often became brown and died in early spring, but did not in a cushion-shaped crown. Needle browning and death is thought to occur by increasing transpiration due to a thinner cuticle or mechanical damage to the cuticle by wind-born snow and ice particles. To confirm whether the needle browning and death in the flagged crown of Abies mariesii Mast., in the alpine region of Japan conform with this concept, we assessed mechanical damage of the needle cuticle in a timberline ecotone and evaluated the effect of cuticle thickness on cuticular resistance. Mechanical damage on needle cuticles of A. mariesii was not observed. In the cushion-shaped crown, epicuticular wax covered the cuticle and plugged stomatal antechambers. In the flagged crown, epicuticular wax was mostly absent. Cuticular resistance in the flagged crown was lower than that in the cushion-shaped crown. However, the cuticle in the flagged crown was thicker than that in the cushion-shaped crown. The needle browning and death in the flagged crown of A. mariesii occurred even though needle cuticles were not mechanically damaged. The thicker cuticle of the flagged crown may play a role in other stresses. To estimate desiccation stress in relation to the cuticle, we need to elucidate not only cuticular resistance and cuticle thickness, but also cuticle quality and structure.

ACID FOG DAMAGE OF BEECH AT HINOKIBORAMARU-TANZAWA, JAPAN

Decline of Japanese beech forest has been obvious at the Tanzawa Mountains near Tokyo. Observations on fog, precipitation and stem flow were performed during June-October, 1994 on both, NE (with healthy beech) and S-SE (with unhealthy beech) slopes, collecting samples biweekly. Chemical species were determined by ion- chromatography and pH meter. The results show that the pH of the precipitation was between 4.4–5.2, with no clear differences in the concentrations of chemical species between both sites and seasons. The pH of the fog water was high from May to the middle of August and lower from mid-August to October at both sites. The concentrations of most chemical species were higher in fog samples than precipitation samples. The concentrations of potassium and magnesium ions in the stem flow of an unhealthy tree on the S-SE slope were higher than those on the NE slope. In June-October, 1995, ten pairs of event fog water samples were obtained with indicating that the pH was below 4.0 for five pairs of event fog water samples. Lower pH value was obtained on the S-SE slope than on the NE slope in every event. Concentrations of most chemical species were always higher in the fog samples from the S-SE slope than those from the NE slope. The equivalent ratio of NO3 −/nss SO4 2− was higher in the fogs of higher acidity. From the result of atmospheric current analysis using a model, it was suggested that the polluted air over the Tokyo Metropolitan Area affected the fog acidity directly and also indirectly, according to the pressure patterns.