Seasonal development of the photosynthetic performance of Norway spruce (Picea abies [L.] Karst.) under magnesium deficiency

In order to investigate the influence of different magnesium nutrition on photosynthesis, one hundred 6-year-old spruce trees derived from one clone were planted in October 1990 into a special out-door experimental construction, where they were cultivated in sand culture with an optimal supply of nutrients, except magnesium, via circulating nutrient solutions. Magnesium was added to the nutrient solutions in three different concentrations, varying from optimal to severe deficient supplies. During the first vegetative period in 1991, photosynthetic performance and carboxylation efficiency were measured under saturating light, controlled CO₂ conditions, optimal temperature and humidity, using a minicuvette system.During summer, the trees under moderate magnesium deficiency developed tip yellowing symptoms on older needles, while the youngest needles remained green with unchanged chlorophyll contents. Trees under severe magnesium deficiency showed yellowing symptoms on all needle age classes combined with decreased chlorophyll contents in the youngest needles as well. In comparison with the controls, the photosynthetic performance of the 1-year-old needles was significantly lower in both deficiency treatments. The same was observed in the youngest needles of the trees under severe deficiency. Trees under moderate deficiency treatment decreased in photosynthetic performance during the summer without reduction of chlorophyll contents. The reduction of photosynthetic rates corresponded to a decrease in carboxylation efficiency, which is taken as a measure of the activity of the enzyme ribulose-1,5-bisphosphate carboxylase. This reduction, together with the observed increase of carbohydrate contents in needles of trees growing under magnesium deficiency, led to the assumption that the photosynthetic carbonfixation is reduced as a consequence of the accumulation of carbohydrates.     

Magnesium deficiency treatment causes reductions in photosynthesis of well-nourished Norway spruce

In order to investigate effects of magnesium deficiency on Norway spruce [Picea abies (L.) Karst.] photosynthesis, 100 well-nourished 5-year-old spruce trees were grown in sand culture, individually supplied with circulating nutrient solutions. Mineral nutrients were added to the nutrient solutions in optimal quantities and optimal relations to nitrogen. Magnesium was supplied at 0.203, 0.041 and 0.005 mM in order to simulate optimal nutrition, moderate deficiency and severe deficiency. Parameters of photosynthetic gas exchange, chlorophyll, magnesium and starch concentrations were determined in current-year and 1-year-old needles during one growing season. By mid May — 6 months after onset of the Mg deficiency treatments in late autumn — CO₂-assimilation rates of 1-year-old needles were significantly decreased independent of the severity of the deficiency treatment, whereas the chlorophyll concentrations did not differ from the controls. The occurrence of yellowing symptoms during July did not further influence the Mg deficiency effect on photosynthesis. In contrast to 1-year-old needles, significant reductions of photosynthesis and chlorophyll in current-year needles were only caused by severely deficient Mg supply. Mg deficiency affected carboxylation efficiency but not light use efficiency. From the accumulation of starch in the needles, up to 30-fold of the controls, the conclusion has been drawn that reactions of CO₂-fixation were affected by reduced carbohydrate export. The light-dependent pigment reduction, leading to the typical tipyellowing of needles, clearly reflects a secondary effect of Mg deficiency.     

Two types of chemolithotrophic nitrification in acid heathland humus

Tree decline has been observed recently in 25–30 year old pine stands inCladina andCalluna-type heath forests in the Hämeenkangas and Pohjankangas areas of southwestern Finland. The trees had grown more or less normally for 15 years. During the following 7 to 8 years increased growth occurred. From 1982 to 1984, however, the trees revealed a sudden reduction in height increment. Additionally, some trees were marked by poor apical shoot dominance. Occasionally complete crown dieback was observed. The trees retained only one to three years' needles. These needles often were characterized by a brownish yellow discoloration.Chemical foliar and soil analysis indicate both a nitrogen deficiency and a deficiency in calcium and magnesium related to the relatively high aluminium levels in the soil. In the needles of affected trees phosphorus and especially potassium concentrations were higher than normally.The low content of nitrogen, calcium and magnesium in the soil is related to the acidic, nutrient-poor bedrock, and the low cation exchange capacity. Also the leaching of nutrients, the shallow and poor quality of the humus layer, and the removal of nutrients by tree harvesting may have effected on the nutritional disturbances.     

Effects of varied soil nitrogen supply on Norway spruce (Picea abies [L.] Karst.)

During a seven-month period the effect of different nitrogen (N) availability in soil on growth and nutrient uptake was studied in three-year-old Norway spruce (Picea abies [L.] Karst.) trees. The plants were grown in pots on N-poor forest soil supplied with various amounts and forms (inorganic and organic) of N. Increasing supply of inorganic N (as NH₄NO₃) increased the formation of new shoots and shoot dry weight. The root/shoot dry weight ratio of new growth was drastically decreased from 1.6 in plants without N supply to 0.5 in plants supplied with high levels of NH₄NO₃. This decrease in root/shoot dry weight ratio was associated with distinct changes in root morphology in favour of shorter and thicker roots. The addition of keratin as organic N source did neither affect growth nor root morphology of the trees. The amount of N taken up by plants was closely related to the supply of inorganic N, and trees supplied with highest levels of NH₄NO₃ also had the highest N contents in the dry matter of needles and roots. In contrast, N contents in needles of trees grown without additional N, or with keratin supply, were in the deficiency range. Supply of NH₄NO₃ decreased the contents of phosphate (P) and potassium (K) and therefore markedly increased N/P and N/K ratios in the needles. On the other hand, the contents of calcium (Ca), magnesium (Mg), and manganese (Mn) in the needles were increased in the plants supplied with inorganic N, suggesting high soil availability and promotion of uptake of these divalent cations by high nitrate uptake. The observed effects on root/shoot dry weight ratio, root morphology, and mineral nutrient composition of the needles indicated that high inorganic N supply may increase above-ground productivity but at the same time decrease the tolerance of trees against soil-borne (e.g. deficiency of other mineral nutrients) stress factors. Deceased 21 September 1996 Deceased 21 September 1996     

Influence of potassium and manganese on growth and uptake of magnesium by soybeans (Glycine max (L.) Merr. cv. Bragg)

Summary Soybean (Glycine max (L) Merr. cv. Bragg) seedlings were grown in nutrient solutions to evaluate the response to manganese nutrition as affected by potassium supply. In solutions containing 275 M manganese, increasing the solution concentration of potassium from 1 mM to 10 mM alleviated symptoms of manganese toxicity, decreased manganese concentrations in the leaves and increased dry matter yields of the plants. The reduction in manganese toxicity was brought about by a reduced rate of root absorption of manganese at high potassium supply levels.Increasing the supply of either potassium or manganese decreased the leaf concentration of magnesium although there were no apparent symptoms of magnesium deficiency in any treatment. The reduced concentration of magnesium in the leaves was due to effects of potassium and manganese on the rate of root absorption of magnesium.Under manganese deficiency conditions, growth was reduced and manganese concentrations in plant parts were very low; there was no effect of potassium supply when manganese was absent from the nutrient solution.     

Seasonal ionic fluctuations and annual growth rates in stands of Pinus silvestris L. and Picea abies Karst. (L.)

Seasonal variations in potassium, calcium, nitrogen, phosphorus, and magnesium contents of needles from two different stands of Pinus silvestris L. and Picea abies (L.) Karst. were determined monthly during a period of two years, and the growth rates once a year during five years. The stands were growing on sand or on clay. The supposed transport of potassium to the shoot during autumn could not be demonstrated. The nutrient status of pine and spruce growing on sand and clay was very similar, except for calcium and magnesium. There were only small differences in ion contents between pine and spruce, except for calcium. On soils with low water-holding capacity, the available water in dry years is the limiting factor affecting the growth rate of spruce but not of pine. It is concluded that the nutrient status of the leaves, the growth rates and the available water in the soil are factors that should have an important part in any discussion of fertilization of coniferous forests.     

Chemical composition and biochemical changes in needles of Scots pine (Pinus sylvestris L.) stands at different stages of decline in Bulgaria

Water content was decreased in the needles of Scots pines (Pinus sylvestris L.) with moderate and slight levels of stress-induced decline. In pines with moderate decline a misbalance of foliar nutritional elements was observed. In 1-year-old needles of afflicted pines, the content of potassium and calcium was decreased, but the content of phosphorus, magnesium, iron, manganese and of the heavy metals cadmium and lead was increased. In 2-year-old needles of damaged pines, the content of calcium, iron and of the heavy metal cadmium was decreased, but the content of phosphorus, magnesium, manganese and zinc was increased. Potassium deficiency and zinc levels above toxicity tolerance characterized the nutrient status of declined Scots pines. In the same pines chlorophyll a/b ratio was diminished in 2-year-old needles and the protein content was increased in 1-year-old needles but was decreased in 2-year-old needles.In the second phase of the study 25- and 40-year-old pines with slight decline were examined. An increase in the protein content of 2-year-old needles of 25-year-old declined pines and of both 1- and 2-year-old needles of older 40-year-old pines was observed. A 280% increase of peroxidase activity in 1-year-old and a 178% increase in 2-year-old needles of 25-year-old damaged pines were shown. In older 40-year-old damaged pines a slight increase only in 1-year-old needles was seen. The observed age-related decrease in peroxidase activity in older 40-year-old symptomatic pines might be responsible for the lower resistance to decline in older pines. Our results show that clear but complex chemical and biochemical changes are observed in the needles of Scots pines in Bulgaria and that these changes hold promise for use as indicators of stress.     

Developmental disorders in buds and needles of mature Norway spruce, Picea abies (L.) Karst., in relation to needle boron concentrations

Dieback of the terminal shoot and consequently bushy growth induced by boron deficiency have been reported widely throughout the world in several tree species. Recently, similar growth damage was documented in half of the young spruce stands in eastern Finland. To clarify the role of B deficiency, the light microscopic structure of emerging buds and of developing and previous-year needles of mature Norway spruce (Picea abies L. Karst.) from damaged (D stand), partly damaged (PD stand) and healthy (H stand) stands were analysed. The samples, on which needle nutrient concentrations were also determined, were taken seven times between early spring (April) and early winter (November). Cell death characterized by precipitation of the cell content, possibly due to the release of tannins after membrane rupture, was seen in the apex of emerging buds, and this led to fatal damage in about half of the buds in the trees from the D stand, where the needle B concentration was well below the deficiency level of 4–5 mg kg⁻¹. Furthermore, an increase in living cells that accumulated tannins in the vacuoles, which is a common stress and/or defense reaction, was found in the primordial shoots of buds and in the differentiating needles in the PD and D stands. The increase in the areas of the central cylinder and of the xylem found in the needles indicate structural plasticity during needle differentiation to drought. The time frame for bud emergence from late May up to mid-September means that an adequate B supply is necessary throughout the summer in order to avoid fatal bud damage and thus bushy growth of the trees.     

Morphology and response of roots of pasture species to phosphorus and nitrogen nutrition

The root morphology of ten temperate pasture species (four annual grasses, four perennial grasses and two annual dicots) was compared and their responses to P and N deficiency were characterised. Root morphologies differed markedly; some species had relatively fine and extensive root systems (Vulpia spp., Holcus lanatus L. and Lolium rigidum Gaudin), whilst others had relatively thick and small root systems (Trifolium subterraneum L. and Phalaris aquatica L.). Most species increased the proportion of dry matter allocated to the root system at low P and N, compared with that at optimal nutrient supply. Most species also decreased root diameter and increased specific root length in response to P deficiency. Only some of the species responded to N deficiency in this way. Root morphology was important for the acquisition of P, a nutrient for which supply to the plant depends on root exploration of soil and on diffusion to the root surface. Species with fine, extensive root systems had low external P requirements for maximum growth and those with thick, small root systems generally had high external P requirements. These intrinsic root characteristics were more important determinants of P requirement than changes in root morphology in response to P deficiency. Species with different N requirements could not be distinguished clearly by their root morphological attributes or their response to N deficiency, presumably because mass flow is relatively more important for N supply to roots in soil.Section editor: H. Lambers     

Organic anion exudation by ectomycorrhizal fungi and Pinus sylvestris in response to nutrient deficiencies

Low molecular weight organic anions (LMWOA) can enhance weathering of mineral grains. We tested the hypothesis that ectomycorrhizal (EcM) fungi and tree seedlings increase their exudation of LMWOA when supply of magnesium, potassium and phosphorus is low to enhance the mobilization of Mg, K and P from mineral grains. Ectomycorrhizal fungi and Pinus sylvestris seedlings were cultured in symbiosis and in isolation on glass beads with nutrient solution or with sand as a rooting medium, with a complete nutrient supply or with Mg, K, P or N in low supply. Concentrations of all dicarboxylic LMWOA in the rooting medium were measured. Nonmycorrhizal seedlings released predominantly malonate. Colonization with Hebeloma longicaudum decreased the amount of organic anions exuded, whereas Paxillus involutus and Piloderma croceum increased the concentration of oxalate but not the total amount of LMWOA. Phosphorus deficiency increased the concentration of LMWOA by nonmycorrhizal and EcM seedlings. Magnesium deficiency increased the concentration of oxalate by nonmycorrhizal and EcM seedlings, but not the concentration of total LMWOA. Paxillus involutus grown in pure culture responded differently to low nutrient supply compared with symbiotic growth. Ectomycorrhizal fungi did not increase the total concentration of LMWOA compared with nonmycorrhizal seedlings but, depending on the fungal species, they affected the type of LMWOA found.     

Phosphorus deficiency induced by nitrogen input in Douglas fir in the Netherlands

Summary A re-examination of earlier NPK fertilization experiments in Douglas fir stands on sandy soils shows the effects of high nitrogen input by air pollution during the last 10–15 years on plant nutrition at these sites. In 1960, experimental plots showed a positive growth reaction to nitrogen, phosphorus, and potassium fertilization. All suffered from severe phosphorus deficiency in 1984, low phosphorus in the needles was invariably accompanied by a high nitrogen content, with all N/P ratios between 20 and 30. The same conclusion emerges from an independent investigation of nutrient status of a selection of Douglas fir stands. Hence, if stand productivity and a balanced nutrient status of the trees is to be maintained, the increase in atmospheric input of nitrogen calls for supplementary fertilization. Given the current N/P ratios in the needles, a positive growth response to phosphorus fertilization is to be expected.     

Changes in phloem export of sucrose in leaves in response to phosphorus, potassium and magnesium deficiency in bean plants

The effect of varied phosphorus (10 and 250 mmol P m^–3potassium (50 and 2010 mmol K m^–3) and magnesium (20 and1000 mmol Mg m^–3 supply on sucrose, reducing sugars, aminoacids, P, K, and Mg in phloem exudate was studied in bean (Phaseolusvulgaris L.) plants over a 12 d growth period in nutrient solution.Phloem exudates were collected from detached primary leavesusing the EDTA-promoted exudation technique. Compared with controlnutrient-sufficient plants, sucrose export in the phloem exudatewas drastically decreased by K deficiency and, particularly,by Mg deficiency, whereas P deficiency either had no effector stimulated sucrose export. In Mg-deficient plants the rateof sucrose export was decreased to 10–20% of the controlplants. There was a close Inverse relationship between phloemexport and leaf concentration of sucrose: higher leaf concentrationsof sucrose were accompanied by lower phloem export of sucrose.In contrast to sucrose, reducing sugars in the exudates werevery low and not affected by P, K and Mg deficiency. The phloemexport of amino acids was strongly depressed by Mg deficiency,but only slightly by P and K deficiency. Resupplying Mg to Mg-deficientplants for 12 h during the dark or light periods rapidly stimulatedsucrose export. After resup ply of Mg for 24 h and 48 h therate of sucrose export was comparable with the rate in the controlplants. The results demonstrate a key role for Mg in phloem loadingand export of photosynthates from source leaves, especiallysucrose. Inhibition of root growth and development of visualsymptoms of chlorosis in Mg-deficient plants are suggested asconsequences of Impaired phloem loading. In agreement with thisin P-deficient plants where phloem loading was not impaired,chlorosis was absent and root growth was maintained at a highlevel. Key words: Bean, carbon partitioning, magnesium nutrition, phloem transport, phosphorus nutrition, potassium nutrition     

The effect of ozone on the yellowing process of magnesium-deficient clonal Norway spruce grown under defined conditions

During two vegetation periods, young clonal spruce trees (Picea abies (L.) Karst.) with sufficient and poor magnesium (Mg) supply were exposed in the environmental chambers of the GSF phytotron to three levels of ozone (daily means: 18-22, 88-130, and 135-190 microg m(-3); 10% reduction at night). Previous year's needles were examined at 4-week intervals with respect to their contents of Mg, Ca, K, Mn, N, P, and chlorophyll (Chl), various parameters of Chl fluorescence, and the stability of the isolated light-harvesting Chl-a/b-protein complex LHC II. The needles of the two nutrition variants contained more than 0.53 or less than 0.27mg Mg g(-1) needle dry matter, respectively. The ratio of variable to maximal Chl-a fluorescence of the dark-adapted needles, Fv/Fm, and the photoinhibitory quenching of Fv after light treatment, SVi.v, were affected by the Mg content of the needles rather than the ozone levels. Changes of the Chl content and the behavior of the LHC II allowed differentiating between a slow process of needle yellowing occurring under Mg deficiency only, and a rapid process of needle yellowing occurring under the combined action of Mg deficiency and ozone pollution. Only the rapid yellowing process was accompanied by destabilization of the LHC II, and the degree of destabilization was correlated with the ozone concentration present in the days before sampling. The results are consistent with observations obtained at a research site in the Central Black Forest (J Plant Physiol 161 (2004) 423).     

Carbon partitioning in Norway spruce: amounts of fructose 2,6-bisphosphate and of intermediates of starch/sucrose synthesis in relation to needle age and degree of needle loss

Summary Intermediates involved in carbon partitioning between starch and sucrose [dihydroxyacetone phosphate + glyceraldehyde 3-phosphate (TP), 3-phosphoglyceric acid, fructose 6-phosphate (F6P), fructose 2,6-bisphosphate (F26BP), in addition to glucose, fructose, sucrose and starch] were analysed in lyophilized needles of Norway spruce (Picea abies L. Karst). Samples were taken from all distinct parts of first and second order branches and the analysed data related to season, needle age, needle position and degree of needle loss (control and class 2 approx. 30%–40% needle loss). Positive and inverse correlations of F26BP, an important regulator of carbon partitioning between starch and sucrose, and F6P or TP existed in all samples. F26BP levels were highest in developing needles and gradually decreased during maturation, which is possibly indicative of changes in the relative sink strength during development (switch from import to export of sucrose). In class 2 needles the amount of F26BP was significantly increased. Together with nearly unaltered levels of sucrose but only slightly decreased amounts of starch the results can be taken as evidence for impaired carbon export in our class 2 samples. The data are discussed with respect to needle development and a possible impact of both air pollutants and mineral deficiency at the location from which the samples were taken.     

Carbon uptake and respiration in above-ground parts of a Larix decidua × leptolepis tree

Summary Shade needles of hybrid larch (Larix decidua × leptolepis) had the same rates of photosynthesis as sun needles per dry weight and nitrogen, and a similar leaf conductance under conditions of light saturation at ambient CO₂ (A_max). However, on an area basis, A_max and specific leaf weight were lower in shade than in sun needles. Stomata of sun needles limited CO₂ uptake at light saturation by about 20%, but under natural conditions of light in the shade crown, shade needles operated in a range of saturating internal CO₂ without stomatal limitation of CO₂ uptake. In both needle types, stomata responded similarly to changes in light, but shade needles were more sensitive to changes in vapor pressure deficit than sun needles. Despite a high photosynthetic capacity, the ambient light conditions reduced the mean daily (in summer) and annual carbon gain of shade needles to less than 50% of that in sun needles. In sun needles, the transpiration per carbon gain was about 220 mol mol^–1 on an annual basis. The carbon budget of branches was determined from the photosynthetic rate, the needle biomass and respiration, the latter of which was (per growth and on a carbon basis) 1.6 mol mol^–1 year^–1 in branch and stem wood. In shade branches carbon gains exceeded carbon costs (growth + respiration) by only a factor of 1.6 compared with 3.5 in sun branches. The carbon balance of sun branches was 5 times higher per needle biomass of a branch or 9 times higher on a branch length basis than shade branches. The shade foliage (including the shaded near-stem sun foliage) only contributed approximately 23% to the total annual carbon gain of the tree.     

Nutrient distributions and bio-cycle characteristics in both natural and artificial Pinus tabulaeformis Carr. forests in hilly loess regions

Pinus tabulaeformis Carr. forest, the dominant community in Ziwuling Mountain lying in the hilly loess region, was studied for its nutrient distributions and bio-cycle characteristics in both natural and artificial forms. The results showed that the changes in the nutrient contents for different components in the same Pinus tabulaeformis Carr. forest stood in the order of needles > branches > bark > roots > bole. The aboveground nutrient elements in needles, branches, bark bole and litterfalls stood in the order of Ca > N > K > Mg > P, but the nutrients stored in the soil stood in the order of Ca > K > Mg > N > P. The accumulated amounts of nutrients increased first and then decreased with the increased age of the forest. The nutrient amounts reached their maximum when the stand was 30 years old, and decreased greatly when it was 50 years old. The 30-year-old artificial Pinus tabulaeformis Carr. forest had the highest annual accumulated amount of nutrients, and different stands stood in the order of II > III > IV > I. Comparatively, annual accumulated nutrients in different components stood in the order of needles > branches > roots > bark > bole. It was also suggested that the amounts of nutrients annually taken in from and retained in the natural Pinus tabulaeformis Carr. forest were significantly higher than those in artificial forests. The coefficients of nutrient use in various Pinus tabulaeformis Carr. stands stood in the order of Ca > Mg > N > K > P, but the nutrient use efficiency (NUE) of the same element decreased with increased age of the forest. There were no differences in the utilization coefficient and the turnover period of nutrients in both natural and artificial matured Pinus tabulaeformis Carr. forests.     

Nutrient distributions and bio-cycle characteristics in both natural and artificial Pinus tabulaeformis Carr. forests in hilly loess regions

Pinus tabulaeformis Carr. forest, the dominant community in Ziwuling Mountain lying in the hilly loess region, was studied for its nutrient distributions and bio-cycle characteristics in both natural and artificial forms. The results showed that the changes in the nutrient contents for different components in the same Pinus tabulaeformis Carr. forest stood in the order of needles > branches > bark > roots > bole. The aboveground nutrient elements in needles, branches, bark bole and litterfalls stood in the order of Ca > N > K > Mg > P, but the nutrients stored in the soil stood in the order of Ca > K > Mg > N > P. The accumulated amounts of nutrients increased first and then decreased with the increased age of the forest. The nutrient amounts reached their maximum when the stand was 30 years old, and decreased greatly when it was 50 years old. The 30-year-old artificial Pinus tabulaeformis Carr. forest had the highest annual accumulated amount of nutrients, and different stands stood in the order of II > III > IV > I. Comparatively, annual accumulated nutrients in different components stood in the order of needles > branches > roots > bark > bole. It was also suggested that the amounts of nutrients annually taken in from and retained in the natural Pinus tabulaeformis Carr. forest were significantly higher than those in artificial forests. The coefficients of nutrient use in various Pinus tabulaeformis Carr. stands stood in the order of Ca > Mg > N > K > P, but the nutrient use efficiency (NUE) of the same element decreased with increased age of the forest. There were no differences in the utilization coefficient and the turnover period of nutrients in both natural and artificial matured Pinus tabulaeformis Carr. forests.     

Diuretic-associated hypomagnesaemia.

Clinically suspected hypomagnesaemia was confirmed in 21 patients over 12 months; all patients had been exposed to either short-term vigorous diuretic treatment or moderate-dosage long-term treatment. Magnesium depletion was compounded by a hospital diet surprisingly low in magnesium, a local soft water supply, and, in some patients, high alcohol intake. Common presenting symptoms included depression, muscle weakness, refractory hypokalaemia, and atrial fibrillation refractory to digoxin treatment. The administration of magnesium supplements resulted in prompt improvement of all symptoms particularly in the case of refractory atrial fibrillation. Chronic low-grade magnesium deficiency from diuretic treatment is more common than published reports suggest. Older patients are at risk, particularly those who have excessive alcohol intake, a diet low in magnesium, or a soft water supply.     

Effects of long-term CO2 enrichment and nutrient availability in Norway spruce. II. Foliar chemistry

 Branches on 30-year-old Norway spruce trees [Picea abies (L.) Karst.] were exposed to ambient (AMB) or ambient plus 350 μmol CO₂ mol^–1 (EL) for 4 years (except winters), using the branch bag technique (BB). The trees were growing on plots with low (control) and high (irrigated-fertilised) availability of soil nutrients. The seasonal variation in foliar macronutrients and non-structural carbohydrates in current and 1-year-old shoots was monitored throughout the treatment period. When the branches were harvested at the end of treatment, macronutrients were analysed in five age classes of foliage. The concentration of all elements, except magnesium, generally increased in AMB, i.e. a ’bag effect’, but decreased as an effect of EL, i.e. a ’CO₂ effect’. At the final harvest K, P, N and S were reduced in young needles by EL, whereas Mg was reduced in older needles on both plots. A change in needle morphology by EL possibly caused a dilution effect in irrigated-fertilised needles, but not in control needles. Reductions in K and Mg are suggested to be an effect of increased phloem transport from the branch, in consequence of higher rates of carbon fixation in EL. Foliage in BBs had higher concentration of Ca, but there was no significant effect of the EL-treatment, indicating that elevated CO₂ had no effect on stomatal conductance. Quinic acid concentration decreased, but shikimic acid concentration increased in BBs, independently of CO₂ treatment. Concentrations of starch and sugars increased in the EL-treatment, but pinitol decreased. Received: 23 October 1998 / Accepted: 1 December 1998     

Effects of root hypoxia and a low P supply on relative growth, carbon dioxide exchange rates and carbon partitioning in Pinus serotina seedlings

The present study was conducted to determine how 10 weeks of root hypoxia and a low P supply altered relative growth, and carbon acquisition and partitioning in a moderately flood tolerant pine. Pond pine (Pinus serotina Michx.) seedlings were grown in continuously flowing solution culture at 5 or 100 μM P, under aerobic or hypoxic solution conditions. Staggered harvests were used to ascertain changes in biomass allocation and relative growth over time. Carbon dioxide exchange rates (CER) were determined by infrared gas analysis, and needles were analyzed for inorganic P (Pi), sucrose, reducing sugars and starch. Although aeration treatment had no significant effect on shoot dry weight or shoot ontogeny, root dry weight of hypoxic seedlings was significantly lower than that of aerobic seedliings after 8 weeks, regardless of the P treatment. Mean relative growth rates (RGR) of roots in the high P treatment initially decreased under hypoxia, but recovered by the sixth week with the production of adventitious roots. Two weeks of hypoxic growth conditions decreased CER and stomatal conductances of seedlings in the high P treatment by more than 30% relative to their aerobic counterparts. Stomatal closure was not accompanied by a decrease in intracellular CO₂, but was accompanied by an increase in starch accumulation. Recoveries of CER, stomatal conductance and carbohydrate metabolism coincided with the recovery of root growth. Low P growth conditions did not significantly affect shoot or root dry weight until the sixth week of treatment. However, differences in seedling RGR, particularly needle RGR, were discernable during the first 2 weeks. Low P treatment effects on CER paralleled changes in needle RGR, with needle RGR more affected than CER. After 6 weeks, CER of aerobically grown seedlings in the low P treatment were only 15% lower than CER of seedlings in the high P treatment, despite a 31% and 75% reduction in needle RGR and Pi concentrations, respectively. Increased starch concentrations of recently expanded needles at this time were probably a result of diminished growth. The inhibitory effect of a low P supply on shoot growth, more specifically on needle expansion and emergence of new fascicular needles, probably limited net carbon fixed per plant more than any direct effect of low P on CER per se.