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Free proline changes in Pinus taeda L. callus in response to drought stress.

The amino acid, proline (PRO), may have an important role in plant adaptation to drought stress. To study the effect of drought stress on PRO content of pine tissues, we measured free PRO by high pressure liquid chromatography in control ( -0.4 MPa) and drought-stressed ( -0.8, -1.0, -1.3, -1.6 MPa) callus tissue of Pinus taeda L. after eight weeks of growth. Drought stress was induced by adding polyethylene glycol (PEG) to the nutrient media. The relation between PRO accumulation and water potential was influenced by composition of the medium and temperature. Callus growing in media with water potentials of -0.8 MPa showed no increase in PRO compared to control callus in media at -0.4 MPa. When callus tissue was subjected to low water potentials (-1.6 MPa), endogenous PRO concentration increased 40-fold, while callus fresh weight decreased by more than 90%. When exogenous PRO was added to the nutrient media, endogenous PRO was directly proportional to the exogenous PRO concentration rather than reduction in callus growth. Thus low water potential in callus results in endogenous PRO accumulation and large reductions in callus fresh weight growth. Proline accumulation in pine cells appears to be related to mechanisms associated with tolerance to desiccation.

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Nutrient retranslocation within the foliage of Pinus sylvestris.

During the period 1983-1987, retranslocation of N, P, K, Ca, Mg, Mn, Zn, Fe, B and Al within the foliage of Scots pine (Pinus sylvestris L.) was studied in sapling, pole-stage and mature stands in eastern Finland. Needle concentrations of the mobile nutrients N, P, and K varied seasonally because of retranslocation. In unfertilized plots, needle contents of N, P and K decreased 62-92% during senescence and needle dry weight decreased 19-51%. In some years, needle contents of Ca, Mn, Zn, Fe, B and Al increased during senescence. Retranslocation was more efficient from needles with greater dry weight. In the fertilized, pole-stage stand, needle dry weight did not decrease at senescence four years after fertilization and nutrient content decreased less than in the other years. In the mature stand, there were no clear differences in needle dry weight and nutrient changes during senescence between years or between unfertilized and fertilized plots.

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Nutrient dynamics in singleleaf pinyon (Pinus monophylla Torr & Frem.) needles.

Nutrient retranslocation from existing foliage of singleleaf pinyon pine (Pinus monophylla Torr. & Frem.) did not meet the nutrient demands of new needles on the same branch. Singleleaf pinyon pines, with an average age of 177 years, located at an elevation of 2300 m on a ridge in The Sweetwater Range in Nevada, USA retained needles for 18 or more years and increased needle number per branch by 17-18% annually. With age, needles gained weight and accumulated Ca, Mg, Fe, Zn, P, and Mn. The amount of N per needle showed no consistent trend with needle age. Only Cu and K declined in concentrations and amounts with needle age and these differences were not statistically significant. Retranslocation of nutrients at needle abscission could provide only 2 to 13% of the N, P, K, Fe, Cu, and Mg required by new needles. We conclude that nutrient capital in existing foliage constitutes a long-term nutrient reserve for the tree rather than a mobile nutrient pool.

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Carbon and phosphorus partitioning in Pinus serotina seedlings growing under hypoxic and low-phosphorus conditions.

Ten-week-old pond pine (Pinus serotina Michx.) seedlings were grown in solution culture at 5 or 100 microM P and under aerobic or hypoxic solution conditions. After 6 and 10 weeks in the treatments, changes in relative growth rate (RGR), P acquisition and allocation, and carbohydrate partitioning were determined by analyzing tissue for total P, soluble sugars and starch. Six weeks of low-P growth conditions decreased seedling dry weight and the ratio of shoot dry weight to root dry weight (S/R) by 39 and 51%, respectively, in comparison to seedlings from the aerobic, high-P (control) treatment. Mean RGRs of shoots in the low-P treatment were reduced by 33%, whereas root growth was unaffected. After 10 weeks of low-P growth conditions, however, both shoot and root RGRs were significantly reduced, and plants had lower S/R ratios than in any other treatment. Slowed shoot growth was accompanied by starch and nonstructural carbohydrate accumulation in needles, indicating that needle growth was not limited by carbohydrate supply. Six weeks of low-P growth conditions decreased total seedling P by 75%, reflecting a 97% reduction in the net uptake rate (NUR). Shoot NUR as a fraction of seedling NUR was also greatly reduced in the low-P treatment, indicating that low-P growth conditions affected P translocation to the shoot more than P accumulation by roots. In contrast, 6 weeks of hypoxic growth conditions decreased total dry weight of seedlings in the high-P treatment by 41% relative to their aerobic counterparts. Root growth was affected more than shoot growth, however, and S/R ratios increased. After 10 weeks, S/R ratios doubled, primarily because of the reduction in root RGR. Nevertheless, roots of hypoxic seedlings contained a higher percentage of total seedling P than their aerobic counterparts. Net P acquisition per seedling decreased by more than 50% under hypoxic growth conditions, as a result of reductions in both root RGR and seedling NUR. Starch accumulation in shoots of hypoxic seedlings reflected reductions both in root growth and in transport of carbohydrates to nonwoody roots. Carbohydrate availability did not appear to be limiting growth of hypoxic woody roots, which are well-aerated internally, but it may have limited metabolic processes in nonwoody roots of seedlings from the high-P treatment.

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Characterization of glutathione S-transferase from dwarf pine needles (Pinus mugo Turra).

Glutathione S-transferase activity conjugating xenobiotics with glutathione (GSH) was found in extracts from needles of dwarf pine (Pinus mugo Turra). In vivo incubation of needle segments with the herbicide fluorodifen at 25 degrees C resulted in conversion of the xenobiotic to water-soluble products at initial rates of 0.7 nmol h(-1) g(fw) (-1). At 15 degrees C, the initial rate of product formation was decreased to 0.1 nmol h(-1) g(fw) (-1). In vitro conjugation studies with chloro-2,4-dinitrobenzene (CDNB) and 1,2-dichloro-4-nitrobenzene (DCNB) as model substrates gave apparent K(m) values of 0.5 mM GSH and 1.14 mM CDNB in the GSH/CDNB system and 0.3 mM GSH and 0.44 mM DCNB in the GSH/DCNB system. The pH optimum was between 7.7 and 7.9 for both the GSH/CDNB and the GSH/DCNB systems. The temperature optimum for these model substrates was between 30 and 35 degrees C, and only minute amounts of enzyme activity were detected at 15 degrees C. The activation energy in the temperature range of 15 to 30 degrees C was 46 kJ mol(-1). Dwarf pine glutathione S-transferase exhibited an approximate molecular weight of 52 kD.

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Mild ozone exposure alters (14)C dynamics in foliage of Pinus taeda L.

To explore the physiological mechanisms underlying ozone-induced growth reductions in loblolly pine (Pinus taeda L.), seedlings were exposed to sub-ambient (charcoal-filtered), ambient or twice-ambient ozone in open-top chambers for three growing seasons. In the final year of exposure, current-year needle fascicles were labeled with (14)CO(2) and the incorporation of (14)C into biochemical fractions was followed for 48 hours. Irrespective of ozone treatment, losses of (14)C-assimilates from foliage to respiration and translocation were minimal during the first 3 hours, whereas more than 60% of the label was lost during the next 45 hours. Radiolabel in sugar decreased rapidly after a lag period, roughly paralleling the pattern of total (14)C loss. The amount of (14)C label in starch and lipids plus pigments remained constant throughout the 48-hour chase period, whereas the amount of (14)C label in other fractions showed a net decrease over the 48-hour chase period. Ozone treatments altered foliar carbon dynamics in two ways: (1) ozone exposure increased foliar (14)C retention up to 21% for the first 5 hours after labeling, but not thereafter, and (2) ozone exposure decreased partitioning of (14)C into starch and increased partitioning of (14)C into organic acids, residue, and lipids plus pigments, indicating an intensified partitioning of carbon to injury and repair processes. Both short-term carbon retention and diversion of carbon from storage compounds to repair processes are foliar mechanisms by which ozone exposure could decrease growth in loblolly pine seedlings.

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Leaf water status and root system water flux of shortleaf pine (Pinus echinata Mill.) seedlings in relation to new root growth after transplanting.

Water relations and root growth of shortleaf pine (Pinus echinata Mill.) were studied four weeks after seedlings from a half-sib family had been transplanted to one of three regimes of soil water availability at a root zone temperature of either 15 or 20 degrees C. About one-third of the variation in new root growth was explained by the root zone environment. The interaction between root zone temperature and soil water availability accounted for 10% of the variation in new root growth. In the most favorable root environment, new roots averaged 620 mm(2) of projected surface area. Leaf water potential increased exponentially with new root projected surface area, becoming constant at about 300 mm(2). Leaf conductance and root system water flux increased linearly with new root growth.

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Root growth and water use efficiency of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and lodgepole pine (Pinus contorta Dougl.) seedlings.

One-year-old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and lodgepole pine (Pinus contorta Dougl.) seedlings were grown for 17 weeks in 100-cm deep, 7.8-liter containers. Two Douglas-fir provenances, one from a wet and one from a dry site in coastal British Columbia, and two lodgepole pine provenances, one from a wet and one from a dry site in interior British Columbia, were grown in wet (522% water content) or dry (318% water content) peat/vermiculite soil in a factorial design. Each container was sealed so that water loss occurred only through the seedling. Five harvests were made at three to five week intervals and water use, dry matter increment, root length and root weight were determined at each harvest. Stomatal conductance and shoot water potentials were measured during the last 12 weeks of the experiment. Lodgepole pine seedlings had greater dry matter production, water use, stomatal conductance and new root length than Douglas-fir seedlings. New root weight of lodgepole pine seedlings exceeded that of Douglas-fir seedlings during the last five weeks of the experiment, and specific root length (root length per unit root weight) of new roots was higher for lodgepole pine seedlings throughout the experiment. Douglas-fir seedlings showed higher water use efficiency (WUE) than lodgepole pine seedlings, and both species showed higher WUE in the dry soil treatment. Douglas-fir seedlings had lower water potentials and higher water uptake rates per unit of new root length than lodgepole pine seedlings, although water uptake rates per unit of root dry weight showed little difference between species. Soil water treatment influenced specific root length of new roots, water uptake per unit of new root length, and WUE in Douglas-fir seedlings more than in lodgepole pine seedlings.

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Analysis of needle proteins and N-terminal amino acid sequences of two photosystem II proteins of western white pine (Pinus monticola D. Don).

A method is described for extracting needle proteins from mature western white pine (Pinus monticola D. Don) trees. Extracted proteins were separated into 26 components by sodium dodecyl sulfate polyacrylamide gel electrophoresis. The predominant protein component (32-39%) had a molecular weight of 57 kDa. The second most predominant protein component (14-25%) had a molecular weight of 16 kDa. The amino acid composition of the twenty-six protein components was determined and a few selected proteins were subjected to further amino acid sequence analysis. Of these, two proteins, designated as Pin m I and Pin m II, were identified by sequence homology with other Photosystem II proteins. The Pin m I and Pin m II proteins had molecular weights of 22.6 kDa and 23.4 kDa, respectively. Pin m I had a much higher proline content than Pin m II.

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Early and late root formation in epicotyl cuttings of Pinus sylvestris after auxin treatment.

Auxin stimulated rooting of epicotyl cuttings of Pinus sylvestris irrespective of the physiological stage of the plant from which the cuttings were taken. The epicotyl cuttings were divided into two groups, those that differentiated roots early (within 6 weeks after cutting) and those that differentiated roots late (more than 6 weeks after cutting). Auxin treatment significantly stimulated the frequency of early rooting of epicotyl cuttings, whereas it had no effect on the frequency of late rooting of epicotyl cuttings. The number of roots per rooted cutting was significantly higher after auxin treatment both on early and late rooted cuttings. Anatomical studies showed that most of the roots on both early and late rooted cuttings developed from wound tissue. However some types of rooting were found only after auxin treatment of early rooted cuttings, e.g., roots that developed from resin duct wound tissue or from vascular tissue or in the pith.

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The effects of acid rain and ozone on biomass and leaf area parameters of shortleaf pine (Pinus echinata Mill.).

Shortleaf pine (Pinus echinata Mill.) seedlings in 24 open-top chambers were exposed to combinations of ozone (carbon-filtered (control), ambient, 1.7 x ambient, and 2.5 x ambient) and acidic precipitation (pH 5.3, 4.3 and 3.3) for 16 months (1989 harvest) or 28 months (1990 harvest). Although the effects of acid rain were generally not significant, there was a trend toward increased aboveground biomass and leaf area in seedlings subjected to the low pH treatments. Because N concentrations in the soils generally increased with decreasing pH, we concluded that the effects of acid rain on aboveground biomass and leaf area were a consequence of an increasing concentration of soil N. In the 1989 harvest, seedlings in the 2.5 x ambient ozone treatment had significantly less biomass in all aboveground plant components and significantly less total leaf area than seedlings in the 1.7 x ambient ozone treatment. In the 1990 harvest, there were no significant effects of ozone on total aboveground biomass, although there was a trend toward reduced biomass in seedlings in the 2.5 x ambient ozone treatment. Both total leaf area and leaf biomass were significantly less in seedlings exposed to 2.5 x ambient ozone for 28 months than in both control seedlings and seedlings in the 1.7 x ambient ozone treatment. The greater, but not always significant, aboveground biomass and leaf area of seedlings in the 1.7 x ambient ozone treatment compared with control seedlings may be associated with the observed increase in soil nitrate concentration as a result of increased rates of leaf senescence and litterfall.

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Vascular cambial sucrose metabolism and growth in loblolly pine (Pinus taeda L.) in relation to transplanting stress.

Sucrose synthase (SS) was the dominant enzyme of sucrose metabolism in both stem and root vascular cambial zone tissues of nursery-grown loblolly pine (Pinus taeda L.) seedlings. Acid invertase (AI) and neutral invertase (NT) activities were generally less than 10% of the SS activity in both tissues. In both cambial tissues, seasonal patterns of enzyme activity were observed for SS but not for AI or NI. The seasonal patterns of SS activity in stem and root cambia paralleled the periodic growth of stems and roots. Stems had high SS activity and growth during summer and early fall. Roots had substantial SS activity and growth during summer and fall, but SS activity and growth were even higher in winter. When seedlings were transplanted, about eight months elapsed before stem and root cambia resumed rates of growth and sucrose metabolism similar to those in control nontransplanted seedlings. Two months after transplanting, root SS was at its lowest, whereas AI activity in transplants was 50% higher than in control nontransplanted seedlings. In stems, SS activity decreased in response to transplanting, whereas AI and NI activities did not change appreciably. In loblolly pine tissues, SS was specific for uridylates, whereas the nucleotide triphosphate-dependent phosphofructokinase (NTP-PFK) had similar activity with either UTP or ATP. Except in winter, the NTP-PFK was less active than the pyrophosphate-dependent phosphofructokinase (PPi-PFK) during all seasons. The PPi-dependent PFK activity in nontransplanted seedlings followed similar seasonal and spatial patterns to those of SS activity. In actively growing tissues, such as stem cambial tissues in summer and root cambial tissues in winter, the measured total PFK to SS ratio ranged between 1.5/1 and 3/1. In contrast, in less actively growing tissues or transplanted seedlings, a greater decrease occurred in SS than in PFK activity, hence the ratio rose to as high as 12/1. It was concluded that: (1) SS was the dominant enzyme for sucrose metabolism in root and stem cambial tissues of loblolly pine seedlings; (2) both SS and PPi-PFK in the cambial tissues can be used as biochemical indicators of growth sink strength in stems and roots; and (3) both enzymes can be used as indicators of seedling stress caused by events such as transplanting and winter freezing.

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The relationship between crown size and ring width in Pinus sylvestris L. stems: dependence on indole-3-acetic acid, carbohydrates and nitrogen in the cambial region.

Indole-3-acetic acid (IAA), carbohydrates, total nitrogen and amino acids in the cambial region and bark were measured at the top (10-year-old internode) and bottom (1.3 m) of the main stem of 50-year-old Pinus sylvestris L. trees, having different rates and longitudinal gradients of annual ring width formation. The trees were sampled during the most active period of wood production (June 23, July 15) and at the end of this period (August 23). Trees with a small crown and relatively slow growth rate (S-trees) were compared with trees in the same stand that had a large crown and fast growth rate (F-trees) as a result of thinning and fertilization. The effect of bottom pruning fast-growing trees (pruned F-trees) was also investigated. The F-trees had greater wood production than the S-trees at both the stem top and bottom. The difference was larger at the stem bottom, thus the relative decrease in ring width down the stem was steeper in the S-trees. The amount of sucrose and IAA per cm(2) in cambial region tissues was higher in F-trees than in S-trees. However, the differences in annual ring width between treatments and within trees were not reflected in the concentrations (expressed per gram fresh weight) of these substances, measured either in differentiating xylem, the cambium plus phloem, or in the cambial region as a whole. The concentrations of total nitrogen and amino acids were slightly higher in the F-trees than in the S-trees. Pruning the F-trees reduced wood production, particularly at the stem bottom, inhibited the springtime increase in starch, and decreased the amounts of sucrose and IAA per cm(2) in the cambial region. However, it was evident that the concentrations of sucrose, IAA, amino acid and nitrogen in the cambial region were not related to ring width. During June and July, the concentrations of these substances were generally higher at the stem bottom in the pruned F-trees than in the F-trees. The results provide evidence that it is the activity of the cambium rather than the availability of carbohydrates that determines the allocation of wood production along the tree stem. The results also indicate that tracheid production is not directly related to the IAA concentration in the cambial region.

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Maturation of maritime pine (Pinus pinaster Ait.) seedlings after exposure to a period of continuous light.

Nine half-sib families of maritime pine (Pinus pinaster Ait.) of known adult performance were grown in continuous light at either 25 degrees C or 25/20 degrees C for 18 weeks. They were then exposed to a dormancy induction period followed by a dormancy release period and then grown for a further 9 weeks in a 16-h photoperiod at a day/night temperature of 25/20 degrees C. Seedlings exhibited great diversity in morphology at the end of the first growth period. The number of morphogenetic cycles varied between one and three and the form of the apical meristem ranged from a typical rosette to an adult-like bud. The type of seedling obtained at the end of the first growth period strongly influenced later growth, independently of the temperature regime. Maturity was proportional to the number of morphogenetic cycles achieved during the first growth period and was characterized by short growth duration, small primary needles and a high degree of fixed growth. The state of the apical meristem that underwent the dormancy period had less influence on the rate of maturation than the number of morphogenetic cycles. The time course of maturation was endogenously controlled and varied among traits. Conspicuous morphological differences were not associated with changes in the relationship between growth components at the phenotypic level. However, there seemed to be a shift in the genetic correlations between growth components after first budset.

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Shoot dynamics of Pinus pumila in relation to altitudinal and wind exposure gradients on the Kiso mountain range, central Japan.

Seasonal shoot dynamics of Japanese mountain stone pine (Pinus pumila Regel.) growing at six sites with different altitudes and slope aspects on the Kiso mountain range in central Japan were investigated. The shoots followed a similar growth pattern at all sites; current shoots elongated rapidly during June and July, and the current needles grew during July and August. Final lengths of the current shoots and needles decreased with increasing altitude. Current-shoot lengths were also shorter at the windward sites than at the leeward sites. At all sites, leaf senescence occurred intensively between mid-August and mid-September. Needle longevity, i.e., the age of the oldest living needle attached to a shoot, increased with increasing altitude. Fascicle density, i.e., the number of current needle fascicles on a unit shoot length, was greater at higher altitudes and windward sites than at lower altitudes and leeward sites. Seasonal dynamics of the P. pumila shoots were characterized by simultaneous replacement of old needles with new needles in the early autumn, thus avoiding any loss of canopy photosynthetic production during the growing season. Increases in needle longevity and fascicle density were associated with declining air temperature and increasing wind exposure. Needle longevity and fascicle density were characteristics of adaptive plasticity in P. pumila that prevent a reduction in growth potential in the stressful conditions of alpine regions.

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Analysis of the relationships among O(3) uptake, conductance, and photosynthesis in needles of Pinus ponderosa.

We studied the effects of O(3) uptake on conductance (g(wv)) and photosynthesis (A) in needles of ponderosa pine (Pinus ponderosa Laws.) seedlings exposed for 70 days to one of three O(3) regimes-Low-O(3) (0.1 micro mol mol(-1) daily peak), High-O(3) (0.2 micro mol mol(-1) daily peak), and Low/High-O(3) (alternating 2 days Low-O(3) and 2 days High-O(3)). Seedlings exposed to charcoal-filtered air served as controls. Total O(3) exposures, expressed as ppm-h (the sum of the average hourly concentration in ppm ( micro mol mol(-1)) over the exposure period), were 77, 135, 105 and 4 for the Low-O(3), High-O(3), Low/High-O(3) and control treatments, respectively. Conductance (g(wv)) declined to about 60% of the value in control seedlings by Day 6 in seedlings in the High-O(3) treatment and by Day 37 in seedlings in the Low/High-O(3) treatment, but g(wv) did not decline at all in seedlings in the Low-O(3) treatment. At the end of the 70-day experiment, cumulative O(3) uptake, calculated from measured g(wv) values and assuming an internal O(3) concentration of zero, was 12.2, 13.5, and 14.7 mmol m(-2) for seedlings in the Low-O(3), Low/High-O(3), and High-O(3) treatments, respectively; however, O(3) uptake was reduced by 0, 24, and 36%, respectively, from that expected if there had been no decline in g(wv). With increasing total O(3) exposure, A declined, but the reduction was not strictly cumulative, i.e., A measured on Days 49 and 70 was similar for a given treatment even though both total O(3) exposure and uptake had increased. At the end of the experiment, A at near saturating CO(2) (1000 micro mol mol(-1)) and saturating photosynthetic photon flux density was reduced by about 25, 40 and 50% in seedlings in the Low-O(3), Low/High-O(3) and High-O(3) treatments, respectively, compared to the control seedlings. The ratio of internal to external CO(2) concentrations, an indicator of relative change in stomatal limitation of A, did not change over time and did not differ among treatments, suggesting that A and g(wv) decreased in parallel. After 40-60 days without O(3), A of seedlings in all O(3) treatments was not significantly different. Our data indicate that O(3)-induced stomatal closure was a result of reduced A and that decreased g(wv) reduced O(3) uptake to a rate that needles of ponderosa pine could tolerate without exhibiting further reductions in gas exchange capacity.

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Growth and nutrition of nonmycorrhizal and mycorrhizal pitch pine (Pinus rigida) seedlings under phosphorus limitation.

The association of ectomycorrhizal fungi with tree roots enhances the acquisition of phosphorus (P) from the soil. In addition to increasing the uptake of H(2)PO(4) (-) (P(i)), mycorrhizal fungi may increase the spectrum of P sources utilized by tree roots by mediating the dissolution of insoluble metallophosphate salts or the hydrolysis of organic P compounds. To investigate the role of ectomycorrhizal fungi in enhancing P acquisition by tree roots, pitch pine (Pinus rigida Mill.) seedlings were grown in sand culture with or without the ectomycorrhizal symbiont Pisolithus tinctorius Coker and Couch under various conditions of P limitation. Compared with nonmycorrhizal seedlings, seedlings inoculated with P. tinctorius exhibited a greater capacity to function under P limitation as evidenced by superior growth and the maintenance of normal foliar ion composition at low P(i) concentrations. Nonmycorrhizal seedlings subjected to P-limiting conditions exhibited depressed K and P and elevated Na concentrations in foliage. The association of P. tinctorius with pitch pine seedling roots maintained foliar K concentrations and prevented the accumulation of Na under P limitation. Nonmycorrhizal seedlings were unable to obtain P from either solid AlPO(4) or inositol hexaphosphate (IHP), whereas seedlings inoculated with P. tinctorius utilized AlPO(4), but not IHP as a P source. Root surface acid phosphatase (APase) activity was depressed in roots infected with the mycorrhizal symbiont and was negatively correlated with seedling growth on all P sources. Root APase activity was negatively correlated with foliar P concentrations in seedlings grown on P(i), but was not correlated with foliar P concentrations in seedlings cultured with AlPO(4) or IHP.

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Heat-shock response of Pinus and Picea seedlings.

The effect of a short period of elevated temperature, or heat shock, on protein synthesis was investigated in 2-day-old seedlings of jack pine (Pinus banksiana Lamb.), loblolly pine (P. taeda L.), lodgepole pine (P. contorta Dougl.), black spruce (Picea mariana (Mill.) BSP), and white spruce (P. glauca (Moench) Voss.). In all species, heat shock led to increased [(35)S]methionine incorporation into heat-shock proteins (hsp's) with molecular masses of 83 and 72 kDA. Heat shock also induced synthesis of several low molecular weight proteins that were absent from control seedlings. The low molecular weight hsp's produced by pine seedlings had molecular masses of 27, 24.6, 20.5 and 17.5 kDa, whereas those produced by spruce seedlings had molecular masses of 27.2, 19.8, 18.4, 17.2 and 16.0 kDa. All of the low molecular weight hsp's showed isoelectric variants. Heat shock led to increased [(35)S]methionine incorporation into a group of low molecular weight hsp's that were also present in control seedlings.

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