Magnesium uptake kinetics in loblolly pine seedlings

Recent studies have suggested that the growth of loblolly pine (Pinus taeda L.) has declined in the southern United States and it has been hypothesized that foliar Mg deficiency may play an important role in the perceived decline. Quantitative nutrient uptake models such as the Barber-Cushman model have been used successfully to investigate nutrient uptake by crop species under a variety of field and experimental conditions and may provide one approach to evaluating this question. However, in order to use this approach it is necessary to develop, for the plant species and nutrient of interest, values for maximal nutrient influx rate at high solution concentrations (I_max), the solution concentration where net influx is 0.5 I_max (K_m), and the nutrient concentration below which influx ceases (C_min). As a first step in evaluating the potential of such an approach, two sets of experiments using established solution nutrient depletion techniques were used to define these values for loblolly pine seedlings 180, 240, 365, and 425 days in age. Observed I_max values for Mg range from 7.90E-8 to 1.29E-7 mol cm^–2 s^–1 with younger seedlings having higher values. Values of K_m for all seedling ages were quite similar ranging from 8.69 to 8.58E-3 mol cm^–3. Most importantly, the results of both experiments indicate that during a growth flush, seedlings will withdraw Mg from solution until the concentration is essentially zero (C_min=0). During non-flush periods uptake rates appear to be greatly reduced. Therefore, efforts to model Mg uptake will need to take these differences as well as seedling age influences into consideration.     

Winter nitrogen fertilization of loblolly pine seedlings

Loblolly pine (Pinus taeda L.) seedlings were nitrogen fertilized during winter in a bare root forest tree nursery located in the coastal plain of the southeastern United States. Total application rates were 0, 50, 100, and 200 kg/N/ha applied in split applications 4 weeks apart in January and February. Seedlings were lifted and outplanted in March, 4 weeks after the second fertilization and measured at 3 and 6 months after outplanting. No seedling morphological differences were encountered at the time of lifting and outplanting although seedling shoot nitrogen content was 28% greater in the highest fertilization treatment compared to the check. Shoot nitrogen concentrations fell after outplanting regardless of treatment, decreasing from an average of 1.51% across all treatments at the time of planting to 0.64% at 6 months after planting. When measured at 6 months after outplanting, seedling dry weight and height growth after planting was shown to increase by 12% and 24%, respectively, for the high nitrogen treatment. This and other studies across a variety of sites have found positive post-outplanting seedling growth response after nutrient loading in the nursery.     

Longleaf and loblolly pine seedlings respond differently to soil compaction,water content,and fertilization

Aims

Longleaf pine (Pinus palustris Mill.) is being restored across the U.S. South for a multitude of ecological and economic reasons, but our understanding of longleaf pine’s response to soil physical conditions is poor. On the contrary, our understanding of loblolly pine (Pinus taeda L.) root and shoot growth response to soil conditions is well established.

Methods

We performed a comparative greenhouse study which modeled root length density, total seedling biomass, and the ratio of aboveground:belowground mass as functions of volumetric water content, bulk density and soil fertility (fertilized or not).

Results

Root length density was about 35 % greater in longleaf pine seedlings compared to loblolly pine seedlings, and was reasonably well modeled (R ²?=?0.54) for longleaf pine by bulk density (linear), volumetric water content (quadratic), soil fertility, and the interactions of bulk density, volumetric water content, species, and soil fertility. The aboveground:belowground mass ratio (ABR) increased at both extremes of water content.

Conclusions

This research indicates that young longleaf pine seedling root systems respond more negatively to extremes of soil physical conditions than loblolly pine, and compacted or dry loamy soils should be ameliorated in addition to normal competition control, especially on soils degraded by past management.     

Root response to CO2 enrichment and nitrogen supply in loblolly pine

This paper examines how elevated CO₂ and nitrogen (N) supply affect plant characteristics of loblolly pine (Pinus taeda L.) with an emphasis on root morphology. Seedlings were grown in greenhouses from seeds during one growing season at two atmospheric CO₂ concentrations (375 and 710 μL L^-1) and two N levels (High and Low). Root morphological characteristics were determined using a scanner and an image analysis program on a Macintosh computer. In the high N treatment, elevated CO₂ increased total plant dry weight by 80% and did not modify root to shoot (R/S) dry weight ratio, and leaf and plant N concentration at the end of the growing season. In the low N treatment, elevated CO₂ increased total dry weight by 60%. Plant and leaf N concentration declined and R/S ratio tended to increase. Nitrogen uptake rate on both a root length and a root dry weight basis was greater at elevated CO₂ in the high N treatment and lower in the low N treatment. We argue that N stress resulting from short exposures to nutrients might help explain the lower N concentrations observed at high CO₂ in other experiments; Nitrogen and CO₂ levels modified root morphology. High N increased the number of secondary lateral roots per length of first order lateral root and high CO₂ increased the length of secondary lateral roots per length of first order lateral root. Number and length of first order lateral roots were not modified by either treatment. Specific root length of main axis, and to a lower degree, of first order laterals, declined at high CO₂, especially at high N. Basal stem diameter and first order root diameters increased at high CO₂, especially at high N. Elevated CO₂ increased the proportion of upper lateral roots within the root system.     

Seasonal ectomycorrhizal fungal biomass development on loblolly pine (Pinus taeda L.) seedlings

Ergosterol, a membrane sterol found in fungi but not in plants, was used to estimate live mycelial biomass in ectomycorrhizae. Loblolly pine (Pinus taeda L.) seeds were sown in April 1993 and grown with standard nursery culture practices. Correlations between total seedling ergosterol and visual assessment of mycorrhizal colonization were high during July and August but low as ectomycorrhizal development continued into the growing season. Percentages of mycelial dry weight over lateral roots decreased from 9% in July to 2.5% in November because seedling lateral root dry weight accumulated faster than mycelial dry weight. Total ergosterol per seedling increased from July through February. As lateral root dry weight ceased to increase during winter months, ectomycorrhizal mycelia became the major carbohydrate sink of pine seedlings. No distinctive seasonal pattern of soil ergosterol content was observed. The impact of ectomycorrhizal fungi on plant carbohydrate source-sink dynamics can be quantitatively estimated with ergosterol analysis but not with conventional visual determination.     

Nutrient uptake by intact and disturbed roots of loblolly pine seedlings

Most measurements of nutrient uptake use either hydroponic systems or soil-grown roots that have been disturbed by excavation. The first objective of this study was to test how root excavation affects nitrate uptake. Rates of NO₃^? uptake by mycorrhizal loblolly pine (Pinus taeda L.) seedlings were measured in intact sand-filled columns, hydroponics, and disturbed sand-filled columns. Total nitrate uptake in intact sand-filled columns was higher than in disturbed columns, indicating that disturbance lowers uptake. Transferring plants from the sand-filled columns to hydroponics had little effect on NO₃^? uptake beyond delaying uptake for an hour. The second objective of this study was to determine whether NH₄⁺, Ca²⁺, Mg²⁺ and K⁺ uptake could be studied using sand-filled columns, since previous studies had tested this method only for nitrate uptake. Uptake rates of NH₄⁺ and K⁺ were positive, while Ca²⁺ and Mg²⁺ uptake rates were negative in intact sand-filled columns, indicating that net efflux may occur even without physical disturbance to the root system. The sand-filled column approach has some limitations, but holds promise for conducting nutrient uptake studies with minimal disturbance to the root system.     

Comparative effects of four mycorrhizal fungi on loblolly pine seedlings growing in a greenhouse in a Piedmont soil

Summary A greenhouse study was conducted to evaluate the effect of ectomycorrhizae on loblolly pine (Pinus taeda L.) growing in a Piedmont soil. Pine seedlings were inoculated with one of four species of fungi (Scleroderma aurantium, Pisolithus tinctorius, Thelophora terrestris, andRhizopogon roseolus). The seedlings were grown in pots containing a Cecil sandy clay loam amended to create a gradient of extractable P ranging from 5.9 to 52.5 g/g. After ten months, all colonized seedlings were significantly larger than control seedlings. However, of the four fungi,Scleroderma aurantium mediated a far superior shoot growth response to increasing levels of soil P; the seedlings were significantly larger than those colonized by any other fungus and also had the largest root systems and greatest degree of mycorrhizal colonization.     

Diurnal changes in allocation and partitioning of recently assimilated carbon in loblolly pine seedlings

We examined diurnal fluctuations in acquisition and partitioning of recently assimilated ¹⁴CO₂, and in subsequent allocation and partitioning to roots of loblolly pine (Pinus taeda L.) seedlings. Nonmycorrhizal seedlings were grown under optimal nutrient conditions in continuously flowin solution culture. Shoots of 15-week-old loblolly pine seedlings were labeled with ¹⁴CO₂ for 30 min at four separate labeling times: 1000, 1200, 1400 and 1600 h. Six whole plant harvests were conducted during a 48 h chase period, i.e. 0, 4, 8 12, 24 and 48 h after the end of the labeling and evacuation periods. Although assimilation of ¹⁴CO₂ was constant between 1000 and 1400 h, there were significant differences in partitioning of ¹⁴C-labeled assimilate in needles of all age classes. The highest percentage of recently assimilated ¹⁴CO₂ in the ethanol-soluble fraction of photosynthesizing tissue was observed near the beginning and end of the photoperiod. Partitioning of ¹⁴C in the ethanol-soluble fraction declined between the 1000 and 1400 h labeling eriods, and was accompanied by an increase in partitioning of recently assimilated ¹⁴CO₂ toward starch and a decrease in respiratory losses. These data suggest that most of the ¹⁴CO₂ assimilated at 1000 h was used to support shoot metabolic activities and possibly restore soluble sugar reserves. Peak starch accumulation in needles during the 1400 h labeling period, concomitant with minimal respiratory loss, indicated that photosynthate production exceeded demand and export out of source leaves. A possible feedback regulation of photosynthesis by starch and/or sugar accumulation may be responsible for the observed decline in assimilation of ¹⁴CO₂ during the 1600 h labeling period. Net accumulation of recently assimilated ¹⁴CO₂ in roots was correlated with assimilation rate of ¹⁴CO₂, but independent of partitioning of recently assimilated carbon in photosynthetic tissue. However, the percentage of total seedling ¹⁴C allocated to roots was essentially the same throughout the 48 h chase, regardless of time of labeling and assimilation rate. The data suggest a strong diurnal regulation of starch and soluble sugars synthesized from recently assimilated carbon in needles of loblolly pine seedlings that was independent of assimilation rate. Allocation and transport of recently assimilated carbon to roots of loblolly pine seedlings were not subject to short-term fluctuations in supply and demand.     

A new hypothesis to explain allocation of dry matter between mycorrhizal fungi and pine seedlings in relation to nutrient supply

Nutrient uptake by forest trees is largely dependent on their associated ectomycorrhizal fungi. The presence of extramatrical mycelium produced by ectomycorrhizal fungi allows trees to exploit a larger soil volume. In this paper the effects of macronutrients on the production of extramatrical mycelium are reviewed. It is concluded that elevated levels of nitrogen and, to some extent, phosphorus strongly inhibit the development of extramatrical mycelium. A deficiency of phosphorus, on the other hand, stimulates ectomycorrhizal development. Low levels of phosphorus may offset the negative influence of nitrogen, indicating that the nitrogen effect is indirect. No other macronutrients have been shown to affect extramatrical mycelium significantly, however, very few studies have been made.To explain reduced ectomycorrhizal development under conditions of high N availability, it has been suggested that the host would allocate less carbohydrate to the mycobiont under such conditions owing to a greater demand for carbon by growing shoots. In the present paper an alternative explanation is suggested: The fungus is forced to take up all available nitrogen and must therefore consume the available carbohydrate in order to assimilate it. The surplus of carbohydrates after nitrogen assimilation can then be used to produce fungal mycelium and fruit bodies. However, the total allocation of host carbohydrate to the mycorrhizal fungus is not reduced at elevated levels of N supply. In contrast with previous theories, the present one proposes that it is the fungus, rather than the host which adjusts its carbon allocation patterns to the N supply.     

Effect of mycorrhiza and nitrate nutrition on nitrate reductase activity in Scots pine seedlings

In vivo nitrate reductase (EC 1.6.6.1) activity was measured in seedlings of Scots pine ( Pinus sylvestris L.) inoculated with Cenococcum geophilum (Sow.) Ferd. & Winge, Paxillus involutus (Batsch:Fr) Fr, Piloderma croceum Erikss, & Hjortst, and Suillus variegatus (Fr.) O. Kuntze. The activity was higher in the mycorrhizal pine roots than was previously found in the fungus symbiont alone, but lower than in the roots of nonmycorrhizal pine seedlings. The differences observed in a previous study between the fungal species under pure culture conditions were not found in the present work for mycorrhiza synthezised with the same fungal species. An increase in the nitrate concentration of the nutrient solution increased the proportion of the nitrate reductase activity in the needles. The mycorrhizal root tips had higher nitrate reductase activity than nonmycorrhizal root tips in the same root system.     

Water relations of pine seedlings in relation to root and shoot growth

The effects of water stress on growth and water relations of loblolly and white pine seedlings were studied during series of drying cycles. As mean soil water potential decreased, growth of roots, needles, and buds decreased. Growth of roots during successive severe drying cycles was not uniform, however. A study of needle and root extension showed that of the total growth of roots for 3 7-day drying cycles, only 6% occurred during the third cycle, while needle extension was uniform for the 3 cycles. The difference in response of needles and roots to drying cycles may be attributed primarily to the effect of water stress on the growing region. When subjected to a severe stress, roots matured toward the tip and became dormant, resulting in less growth during subsequent drying cycles. The intercalary growing region of needles, however, was not altered seriously enough by the stress to cause a difference in amount of growth during each drying cycle.

Transpiration of loblolly pine was lower in the second drying cycle than in the first. Needle water potential after rewatering was as high as that of control plants watered daily; root resistance was apparently not important in restricting transpiration during a second drying cycle. Needle diffusion resistance of loblolly pine, measured with a low-resistance diffusion porometer, was slightly higher during the second drying cycle than during the first. In addition, many primary needles were killed during the first period of stress. These factors contributed to the reduction of transpiration during the second drying cycle. Diffusion resistance of Coleus increased and transpiration ceased during the first drying cycle while water potential remained relatively high. After rewatering, both leaf resistance and transpiration returned to the control level, presumably because the stress during the first period of drying was not severe. The diffusion resistances observed for well-watered plants were 30 to 50 sec·cm⁻¹ for loblolly pine, 3 to 5 sec·cm⁻¹ for Coleus, and 4 to 6 sec·cm⁻¹ for tomato. These values agree closely with those reported by other workers.

     

Seasonal development of loblolly pine lateral roots in response to stand density and fertilization

In 1989, two levels each of stand density and fertilization treatments were factorially established in a 9-year-old loblolly pine plantation on a P-deficient Gulf Coastal Plain site in Rapides Parish, Louisiana, USA. In 1995, a second thinning was conducted on the previously thinned plots and fertilizer was re-applied to the previously fertilized plots. The morphology of new long lateral roots was evaluated at 2-week intervals in five Plexiglas rhizotrons per plot of two replications. The overall objective of this study was to evaluate the seasonal initiation of six morphological categories of long lateral roots ( 2.5 cm in length) in response to stand density and fertilization. Lateral root development exhibited a seasonal pattern with the initiation of branched lateral roots predominantly occurring in spring and summer. The initiation of non-branched lateral roots occurred throughout the year regardless of season. Stand density did not affect lateral root morphological development. However, fertilization stimulated the initiation of branched lateral roots that were greater than 1 mm in diameter.     

The effect of soil moisture tension and nitrogen supply on nitrate reduction and accumulation in wheat seedlings

Summary The effect of soil moisture tension on nitrate reductase and on nitrate accumulation in wheat plants was studied. Nitrate reductase activity was inhibited when soil moisture tension was increased to about 3.0 bars associated with a drop in leaf relative water content to about 90 per cent. The decrease in nitrate reductase activity did not result in nitrate accumulation in short-term experiments (10 days) when plants were exposed to only 1–2 cycles of elevated soil moisture tensions. However, when the period of different moisture regimes was extended up to the flag-leaf stage, nitrate accumulated in stressed plants.Significant increase in plant nitrate concentration as a result of increased moisture tensions was only found at the high levels of added nitrogen. On the other hand, moisture tensions had no effect on the content of total nitrogen in wheat shoots, implying that nitrate reduction was rather limiting under stress conditions.An effect of soil moisture tension and nitrogen nutrition on dry matter production by wheat seedlings was also found in the long-term experiment. At the highest dose of soil nitrogen an increase in maximal soil moisture tension from 0.1 to 0.33 bars reduced plant growth; at intermediate nitrogen doses only tension higher than 2 bars reduced growth. Under complete nitrogen deficiency, plant dry matter production was very low and was not affected by soil moisture tensions.Contribution from the Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel. 1972 Series, No. 2185-E.Contribution from the Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel. 1972 Series, No. 2185-E.     

Growth and nutrition of birch seedlings in relation to potassium supply rate

Summary Growth of hydroponically cultivated birch seedlings (Betula pendula Roth.) at sub- and supra-optimum potassium supply rates was investigated. Potassium was supplied either as a relative addition rate (r _k = 5, 10, 15 and 20% increase day^-1) or as fixed concentrations (0.2, 3, 6, 12 and 15 mM) in the culture solution. After an acclimation period the growth rate of the seedlings in the suboptimum treatments reached values close to the treatment variable, the relative rate of K-addition. Deficiency symptoms, in the form of chlorosis and necroses along the leaf margins, developed initially in all suboptimum treatments, but very few new symptoms appeared once the seedlings had reached the phase of steady-state nutrition and growth. At supra-optimum K-supply levels, i.e. at 0.2–15 mM K in the culture solution, no symptoms of deficiency or toxicity developed, and the relative growth rate of the seedlings remained maximum. The relative growth rate of the seedlings was linearly related to the plant K-status for K contents ranging from 0.2 to 1.0% of dry weight (DW). At higher internal K-concentrations, 1.0–3.0% DW, no further increase in relative growth rate was achieved. A shortage of K resulted in a decrease in the net assimilation rate. This effect was counterbalanced by the absence of shift in he leaf weight ratio as well as by the production of relatively thin leaves. The fraction of dry matter allocated to roots decreased in K-limited plants, as did the leaf contents of soluble carbohydrates and starch.     

Interactive effects of soil nitrogen and atmospheric carbon dioxide on root/rhizosphere carbon dioxide efflux from loblolly and ponderosa pine seedlings

We measured CO₂ efflux from intact root/rhizosphere systems of 155 day old loblolly (Pinus taeda L.) and ponderosa (Pinus ponderosa Dougl. ex Laws.) pine seedlings in order to study the effects of elevated atmospheric CO₂ on the below-ground carbon balance of coniferous tree seedlings. Seedlings were grown in sterilized sand culture, watered daily with either 1, 3.5 or 7 mt M NH ₄ ⁺ , and maintained in an atmosphere of either 35 or 70 Pa CO₂. Carbon dioxide efflux (mol CO₂ plant^–1 s^–1) from the root/rhizosphere system of both species significantly increased when seedlings were grown in elevated CO₂, primarily due to large increases in root mass. Specific CO₂ efflux (mol CO₂ g root^–1 s^–1) responded to CO₂ only under conditions of adequate soil nitrogen availability (3.5 mt M). Under these conditions, CO₂ efflux rates from loblolly pine increased 70% from 0.0089 to 0.0151 mol g^–1 s^–1 with elevated CO₂ while ponderosa pine responded with a 59% decrease, from 0.0187 to 0.0077 mol g^–1 s^–1. Although below ground CO₂ efflux from seedlings grown in either sub-optimal (1 mt M) or supra-optimal (7 mt M) nitrogen availability did not respond to CO₂, there was a significant nitrogen treatment effect. Seedlings grown in supra-optimal soil nitrogen had significantly increased specific CO₂ efflux rates, and significantly lower total biomass compared to either of the other two nitrogen treatments. These results indicate that carbon losses from the root/rhizosphere systems are responsive to environmental resource availability, that the magnitude and direction of these responses are species dependent, and may lead to significantly different effects on whole plant carbon balance of these two forest tree species.