ALBERT

Notes: Higher rates of nitrate assimilation are required to support faster growth in enhanced carbon dioxide. To investigate how this is achieved, tobacco plants were grown on high nitrate and high light in ambient and enhanced (700 μmol mol–1) carbon dioxide. Surprisingly, enhanced carbon dioxide did not increase leaf nitrate reductase (NR) activity in the middle of the photoperiod. Possible reasons for this anomalous result were investigated. (a) Measurements of biomass, nitrate, amino acids and glutamine in plants fertilized once and twice daily with 12 mol m–3 nitrate showed that enhanced carbon dioxide did not lead to a nitrate limitation in these plants. (b) Enhanced carbon dioxide modified the diurnal regulation of NR activity in source leaves. The transcript for nia declined during the light period in a similar manner in ambient and enhanced carbon dioxide. The decline of the transcript correlated with a decrease of nitrate in the leaf, and was temporarily reversed after re-irrigating with nitrate in the second part of the photoperiod. The decline of the transcript was not correlated with changes of sugars or glutamine. NR activity and protein decline in the second part of the photoperiod, and NR is inactivated in the dark in ambient carbon dioxide. The decline of NR activity was smaller and dark inactivation was partially reversed in enhanced carbon dioxide, indicating that post-transcriptional or post-translational regulation of NR has been modified. The increased activation and stability of NR in enhanced carbon dioxide was correlated with higher sugars and lower glutamine in the leaves. (c) Enhanced carbon dioxide led to increased levels of the minor amino acids in leaves. (d) Enhanced carbon dioxide led to a large decrease of glycine and a small decrease of serine in leaves of mature plants. The glycine:serine ratio decreased in source leaves of older plants and seedlings. The consequences of a lower rate of photorespiration for the levels of glutamine and the regulation of nitrogen metabolism are discussed. (e) Enhanced carbon dioxide also modified the diurnal regulation of NR in roots. The nia transcript increased after nitrate fertilization in the early and the second part of the photoperiod. The response of the transcript was not accentuated in enhanced carbon dioxide. NR activity declined slightly during the photoperiod in ambient carbon dioxide, whereas it increased 2-fold in enhanced carbon dioxide. The increase of root NR activity in enhanced carbon dioxide was preceded by a transient increase of sugars, and was followed by a decline of sugars, a faster decrease of nitrate than in ambient carbon dioxide, and an increase of nitrite in the roots. (f) To interpret the physiological significance of these changes in nitrate metabolism, they were compared with the current growth rate of the plants. (g) In 4–5-week-old plants, the current rate of growth was similar in ambient and enhanced carbon dioxide (≈ 0·4 g–1 d–1). Enhanced carbon dioxide only led to small changes of NR activity, nitrate decreased, and overall amino acids were not significantly increased. (h) Young seedlings had a high growth rate (0·5 g–1 d–1) in ambient carbon dioxide, that was increased by another 20% in enhanced carbon dioxide. Enhanced carbon dioxide led to larger increases of NR activity and NR activation, a 2–3-fold increase of glutamine, a 50% increase of glutamate, and a 2–3-fold increase in minor amino acids. It also led to a higher nitrate level. It is argued that enhanced carbon dioxide leads to a very effective stimulation of nitrate uptake, nitrate assimilation and amino acid synthesis in seedlings. This will play an important role in allowing faster growth rates in enhanced carbon dioxide at this stage.

Notes: Abstract. Wild radish plants deprived of, and continuously supplied with solution NO−3 for 7 d following 3 weeks growth at high NO−3 supply were compared in terms of changes in dry weight, leaf area, photosynthesis and the partitioning of carbon and nitrogen (NH2-N and NO−3-N) among individual organs. Initial levels of NO−3-N accounted for 25% of total plant N. Following termination of NO−3 supply, whole plant dry weight growth was not significantly reduced for 3 d, during which time plant NH2-N concentration declined by about 25% relative to NO−3-supplied plants, and endogenous NO−3-N content was reduced to nearly zero. Older leaves lost NO−3 and NH2-N, and roots and young leaves gained NH2-N in response to N stress. Relative growth rate declined due both to decreased net assimilation rate and a decrease in leaf area ratio. A rapid increase in specific leaf weight was indicative of a greater sensitivity to N stress of leaf expansion compared to carbon gain. In response to N stress, photosynthesis per unit leaf area was more severely inhibited in older leaves, whereas weight-based rates were equally inhibited among all leaf ages. Net photosynthesis was strongly correlated with leaf NH2-N concentration, and the relationship was not significantly different for leaves of NO3−-supplied compared to NO−3-deprived plants. Simulations of the time course of NO−3 depletion for plants of various NH2-N and NO−3 compositions and relative growth rates indicated that environmental conditions may influence the importance of NO−3 accumulation as a buffer against fluctuations in the N supply to demand ratio.

Notes: Summary The water relations of Picea abies in a healthy stand with green trees only and a declining stand with trees showing different stages of needle yellowing were investigated in northern Bavaria. The present study is based on observations of trees differing in their nutritional status but apparently green on both sites in order to identify changes in the response pattern which might be caused by atmospheric concentrations of air pollutants and could lead to the phenomenon of decline. Transpiration was measured as water flow through the hydroactive xylem using an equilibrium mass-flow measurement system. Total tree transpiration was monitored diurnally, from July 1985 until October 1985 at both sites. The relationship between transpiration and meteorological measurements indicated that transpiration was a linear function of the vapor pressure deficit. No differences in transpiration of green trees were observed between the two sites. Canopy transpiration was 57%–68% of total throughfall and 41%–54% of total rainfall. Due to this positive water balance, soil water potential at 10 and 20 cm depths remained close to-0.02 MPa (max.-0.09 MPa) for most of the summer. Soil water potential was correlated with the difference between the weekly precipitation and transpiration. No differences in the water relations of apparently healthy trees in the two P. abies stands were observed. It is concluded that differences between green trees at the two sites in terms of nutrient relations or growth rate cannot be explained by changes in whole-tree transpiration or soil water status.

Notes: Summary CO2 assimilation rate (A) and leaf conductance (g) were measured in the field on intact branches of 35-year-old Picea abies (L.) Karst. trees, in five plots each in a healthy and a declining stand. The declining site included trees with yellow needles. In order to separate atmospheric effects on gas exchange from effects of nutrient deficiency, direct effects of atmospheric pollutants were studied on green needles of different age classes in plots of trees at different stages of visible decline. The effects of nutrient deficiency on gas exchange were studied on a different group of trees showing needles of various degrees of yellowing. CO2 assimilation of green needles at the same leaf conductance fell somewhat only when needles had reached 5 years of age, the oldest age examined in this study. Leaf conductance decreased with increasing needle age, but green needles in the declining stand had leaf conductances similar to those of needles in the healthy stand. Stomata of needles with different magnesium concentrations responded to light and air humidity in all age classes. Thus, as long as needles were green, no dese effect was detectable up to 5 years of exposure to atmospheric emissions. Since all needles, green and yellow, were exposed to the same pollution levels, differences in gas exchange between green and yellow needles could not be explained simply in terms of long-term direct effects of air pollution. Needle magnesium contents were correlated with needle yellowing. Neither needle color change nor the magnesium concentration were related to g, but CO2 uptake at ambient CO2 levels declined with lower magnesium concentration and greater degrees of needle yellowing.

Notes: Summary The nutrient relations (nitrogen, magnesium, calcium, potassium, and manganese) of the xylem sap of spruce trees, Picea abies (L.) Karst., growing at a healthy and a declining site in Northern Bavaria, were followed on a diurnal and seasonal basis between April and October 1985. There were significant differences between the two sites in the xylem sap concentrations of all elements investigated except nitrogen. Nutrient concentrations remained constant diurnally despite changes in transpiration and xylem water potential. However, during periods between precipitation events, concentrations of elements in xylem sap decreased with decreasing xylem water potential. Apparent differences in needle chlorosis of spruce trees at the two sites were associated with consistent differences in nutrient contents of their xylem sap and needles.

Notes: Summary The development of root tips and apparent ectomycorrhizas was compared in the Fichtelgebirge (FRG) over one growing season in two 30-year-old Picea abies stands, both on soils derived from phyllite but showing varying symptoms of decline. Visual symptoms of tree decline reflected a lower relative and absolute mycorrhizal frequency, a lower number of ectomycorrhizas per m2 leaf area and an uneven vertical distribution of root tips and ectomycorrhizas. The number of apparent ectomycorrhizas per ground area was correlated with the amount of magnesium, calcium, and ammonium, and the pH in the free-drainage soil solution, and with the molar calcium to aluminium ratio in mineral soil extracts. The foliage concentrations of magnesium and calcium were correlated with the numbers of apparent ectomycorrhizas per m2 leaf or ground area. These observations were used to formulate testable hypotheses concerning the role of the root system and the soil environment in forest decline.

Notes: Summary A declining, closed-canopy Picea abies (L.) Karst. stand produced as much crown biomass as a healthy stand, although some trees were chlorotic due to magnesium deficiency. The production of wood per unit of leaf area in both stands was related to the foliar magnesium concentration. Although leaf area index and climate were similar at both sites, stemwood production was 35% lower in the declining than in the healthy stand. Nutritional disharmony, rather than a deficiency in a single element, was identified as the mechanism for reduced tree vigor. The role of nutrient stress in forest decline was detected by partitioning the season into three periods reflecting different phenological stages: a canopy growth period in spring, a stem growth period in summer, and a recharge period during the non-growing season. Needle growth was associated with nitrogen supply. Most of the magnesium supply required to meet the demand for foliage growth was retranslocated from mature needles. Magnesium retranslocation was related to concentration of nitrogen and magnesium in those needles before bud break. Retranslocation from mature needles during the phase of canopy production resulted in chlorosis in initially green needles if the magnesium concentration before bud break was low. Nitrogen concentration in 0-year-old needles generally remained constant with increasing supply, indicating that foliage growth was restricted by the supply of nitrogen. In contrast, magnesium concentration generally increased with supply, indicating that magnesium supply for needle growth was sufficient. Much of the magnesium required for wood production was taken up from the soil because stored magnesium was largely used for canopy growth. Uptake at the declining site was probably limited because of restricted root expansion and lower soil magnesium compared to the healthy site. For this reason only wood growth was reduced at the declining site. Because the recharge of magnesium during the non-growing period is dependent on uptake from the soil, it was more limited at the declining that at the healthy stand. However, as nitrogen uptake from the atmosphere may account for an appreciable proportion of the total uptake, and as its supply in the soil at both sites was similar, an unbalanced recharge of nitrogen and magnesium may have occurred at the declining site. If mature needles are unable to recharge with magnesium in proportion to the uptake of nitrogen, chlorosis is likely to occur during the next canopy growth period.

Notes: Abstract Downward transport of water in roots, in the following termed “inverse hydraulic lift,” has previously been shown with heat flux techniques. But water flow into deeper soil layers was demonstrated in this study for the first time when investigating several perennial grass species of the Kalahari Desert under field conditions. Deuterium labelling was used to show that water acquired by roots from moist sand in the upper profile was transported through the root system to roots deeper in the profile and released into the dry sand at these depths. Inverse hydraulic lift may serve as an important mechanism to facilitate root growth through the dry soil layers underlaying the upper profile where precipitation penetrates. This may allow roots to reach deep sources of moisture in water-limited ecosystems such as the Kalahari Desert.

Notes: Summary This is the first in a series of papers on the growth, photosynthetic rate, water and nutrient relations, root distribution and mycorrhizal frequency of two Norway spruce forests at different stages of decline. One of the stands was composed of green trees only while the other included trees ranging in appearance from full green crowns to thin crowns with yellow needles. In this paper we compare the growth and carbohydrate relations of the two stands and examine relationships among growth variables in ten plots. The declining stand produced 65 percent of the wood per ground area compared with the stand in which all trees were green because its foliage produced less wood at any level of leaf area index. The difference in foliage efficiency between the sites could not be explained by differeneces in climate, competition or stand structure. The declining stand appeared to have lower carbon gain as indicated by a smaller increase in reserve carbohydrates before bud break, and weaker sinks for carbohydrates as indicated by less use of the stored carbohydrates than the healthy stand. Thus, growth reduction was probably related to factors which affect both photosynthesis and, even more, the sinks for carbohydrate.