Growth and mineral nutrient analysis of teak (<Emphasis Type="Italic">Tectona grandis</Emphasis>) grown on acidic soils in south China

Teak (Tectona grandis L.f.) is widely planted in the world due to its high market demand, economic, ecological and social value. Its plantations have mostly been established and expanded into sites that are acidic to severely acidic in southern China. But, there are no available and specific evidence-based nutrient management techniques. To better recognize and understand the relationship between teak tree growth and nutrient content in the foliage and soil and establish nutrient norms are critical to optimally manage these young plantations. We studied the foliar nutrient and soil chemistry in 19 representative teak plantations aged 5–8 years. Regression analysis indicated that the mean annual increment of teak volume was linearly and positively correlated with foliar N, Ca, Fe and B concentrations, with soil base saturation percentage, available P and Zn concentrations, and negatively correlated with soil Al concentration. Only if the Ca and Mg contents in soil were enhanced, could the increase in soil base saturation percentage benefit teak growth. A revised classification of low-and high-yielding stands was established by using a sorting method of principal components over 6 foliar macro and 8 micro elements in a Diagnosis and Recommendation Integrated System (DRIS). Specific DRIS norms for teak plantations in acid soils were derived. The nutrient balance of N, P, K Ca, Mg, Zn, B with Fe or Al, Ca with Mg, and Fe with Al provided a key to promote the growth of teak in acid soils. Meanwhile, soil Zn was also found as a primary trace element that affected teak growth in this study.     

Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Effects of vegetation control on nutrient removal and Fagus orientalis, Lipsky regeneration in the western Black Sea Region of Turkey

Dense Rhododendron ponticum (L.) understories of eastern beech (Fagus orientalis Lipsky) stands in the Black Sea Region (BSR) of Turkey create challenging forest vegetation management problems relative to beech regeneration. Rhododendron traditionally is controlled in Turkish forests with grubbing and bulldozing. The effects of these practices on nutrient removal and natural beech regeneration have not been quantitatively studied. Two woody vegetation control treatments (bulldozing and hand-grubbing) were installed during late summer, 2002 in three mature beech stands with dense rhododendron understories in the Düzce Forest Management Directorate, in the Turkish western BSR. Aboveground biomass of each vegetation component, total aboveground vegetation biomass, nutrient concentrations, organic matter (OM) removal, and total amount of OM nutrients were determined for each woody vegetation control treatment. Soil bulk density and nutrient content, and beech seedling biomass, nutrient content, and natural regeneration also were studied. One year after treatment, the machine site preparation by bulldozing (MSP) that removed understory vegetation and attached roots, reduced mean forest floor OM content by about 84%, when compared to hand-grubbing. Mean soil C, N, K and Mg concentrations on the bulldozed sites were 36, 27, 50 and 55% less, respectively, than those on the grubbed sites. Total C and Mg amounts at the 5–10 cm soil depth were 24 and 47% lower, respectively, for mechanical site preparation (MSP) sites, when compared to grubbed sites. Overall, soil bulk density did not differ significantly between the grubbing and MSP treatments. Frequent passes on designated transects on MSP sites resulted in a significant (P < 0.01) one-third increase in bulk density of the first 20 cm soil depth, when compared to grubbed sites. No natural beech regeneration occurred on untreated control sites. Mineral soil exposure on the MSP sites increased beech seedling germination substantially. One year after treatment, the mean number of naturally regenerated seedlings for MSP sites was 2.5 times greater than for grubbed sites. Woody control treatments had significant effects upon beech seedling chemistry, with N, P, K, Ca and Mg concentrations averaging 35, 47, 12, 33 and 25% lower, respectively, for MSP sites, when compared to grubbed sites. However, mean seedling biomass and nutrient content were significantly greater (P < 0.05) on MSP sites. Long-term effects from windrowing on MSP site residues, associated topsoil and nutrients need to be evaluated.     

Interactive Influences of Fertility and Nodulation on Growth,Nutrition, and Water Relations of European Alder in a Mine Soil

Abstract

Actinorhizal nodulation of European alder (Alnus glutinosa[L.] Gaertn.) as influenced by substrate fertility was assessed for its effects on seedling growth and physiology. After a brief period in seed flats, seedlings were transplanted to a mine soil where they resided for 18 months, and three fertility regimes were imposed by application of differing nutrient solution concentrations while two distinct levels of nodulation were created through methyl bromide fumigation. Ultimately, seedlings in nonfumigated soil had > 5Xthe nodules of those grown in fumigated soil, and rising fertility stimulated nodule formation in the former. Heavy nodulation resulted in a 35% increase in shoot mass and a 60% increase in root mass on average, responses that exceeded those to fertilization for the latter. Seedlings with abundant nodules had higher foliar concentrations of total N, NO₃, K, and Cu and lower concentrations of Ca, Mg, S, Fe, Mn, Zn, Mo, and Al overall, than were found in those with minimal nodulation. In turn, soil in which heavily nodulated seedlings were grown had higher total N, NO₃, Ca, Mg, S, Fe, Mn, Zn, and Al concentrations but a lower Mo concentration than fumigated soil. Fertility regime effects on nutrition were readily apparent, although considerably more so in the mine soil than in foliage. During a simulated drought episode, predawn and midday xylem pressure potentials within the high fertility treatment were lower in heavily nodulated seedlings than in those with few nodules, and the water potentials in soil in which the heavily nodulated seedlings resided were lower overall as well. Collectively, these results provide insight into approaches to most effectively exploit the benefits European alder offers as a reforestation species for difficult sites.     

Responses of two closely related oak species, Quercus robur and Q. petraea, to excess manganese concentrations in the rooting medium

Quercus robur (L.) and Q. petraea (Matt.) Liebl. are European oak species that often grow in forest soils with high soluble manganese (Mn2+) concentrations. We tested the effects of Mn2+ at concentrations of 0.0024 mM (control), 0.24 mM (typical of acidic forest soils) and 1.2 mM (typical of forest soils under strongly reducing conditions) on the growth, tissue anatomy, foliar element concentrations, subcellular element distribution and gas exchange of solution-cultured seedlings. At the highest Mn2+ concentration, seedlings were grown with and without an elevated concentration (1.2 mM) of magnesium (Mg2+). At 0.24 mM Mn2+, foliar Mn concentrations were higher than observed in the field. Vacuoles of the leaf epidermis and mesophyll were the main sites of manganese accumulation. High nutrient solution Mn2+ concentration significantly lowered foliar iron (Fe) and Mg concentrations. Elevated Mg2+ concentration raised the foliar Mg concentrations to control values, but Fe concentrations and gas exchange remained depressed. In seedlings grown in the 1.2 mM Mn2+ treatment without elevated Mg2+ damage to the phloem of the petioles and a reduction in root mass were observed in both species. The effects on shoot and root growth were greatest in Q. petraea. Alleviation of manganese toxicity symptoms by Mg2+ in Q. petraea was less effective than in Q. robur. Our results suggest that the soil solution Mn2+ concentrations that occur in European oak forests are unlikely to affect the distribution and performance of Q. robur and Q. petraea in the field.     

Growth and nutrient use efficiency of water tupelo seedlings in flooded and well-drained soil

Growth characteristics and nutrient utilization rate of Nyssa aquatica L. seedlings grown in pots containing flooded or well-drained soil were compared in a greenhouse study. For most of the growing season, relative height and diameter growth rates, and biomass accumulation rates were greater for seedlings in flooded soil than for seedlings in well-drained soil. The concentration of Fe in the roots of seedlings in flooded soil was almost tenfold greater than that of seedlings in well-drained soil. However, flooding had no effect on foliar Fe concentrations. The flooding treatment resulted in decreased concentrations of N in all component parts and increased concentrations of P in the roots and stem, but it had no effect on foliar P concentrations. In response to flooding, foliar K concentrations decreased, whereas the concentration of K in the roots increased. Flooding had no effect on the K concentration of the stem. Seedlings in flooded soil produced more total biomass per milligram of nutrient absorbed than seedlings in well-drained soil, suggesting that N. aquatica seedlings are more efficient at producing biomass and height growth under hydric conditions than under mesic conditions.     

Relationships between soil and foliar nutrients in young densely planted mini-plots of Pinus radiata and Cupressus lusitanica

The long-term nature of forest crop rotations makes it difficult to determine impacts of forestry on soil nutrients that might be depleted by forest growth. We used small scale, highly stocked plots to compress the length of the rotation and rapidly induce nutrient depletion. In the study, two species (Pinus radiata D. Don and Cupressus lusitanica Miller) are compared under two disturbance regimes (soil undisturbed and compacted), and two fertiliser treatments (nil and plus fertiliser), applied in factorial combination at 33 sites, covering the range of climatic and edaphic variation found in plantation forests across New Zealand. To assess our ability to rapidly highlight important soil properties, foliar nutrient concentrations were determined 20 months after planting. It was hypothesised that the densely planted plots, even at a young age, would create sufficient pressure on nutrient resources to allow development of relationships between properties used as indicies of soil nutrient availability and foliar nutrient concentrations. For both species significant relationships between foliar nutrients and 0–10 cm layer soil properties from unfertilised plots were evident for N (total and mineralisable N) and P (total, acid extractable, organic, Bray-2 and Olsen P). With the exception of Ca in C. lusitanica, foliar K, Ca and Mg were correlated with their respective soil exchangeable cation measures. The results thus confirm the utility of the experimental approach and the relevance of the measured soil properties for forest productivity.

In unfertilised plots foliar N and P concentrations in P. radiata exceeded those in C. lusitanica, the differences being eliminated by fertiliser application. Foliar N/P ratios in P. radiata also exceeded those in C. lusitanica. In contrast to N and P, foliar K, Ca and Mg concentrations were all higher in C. lusitanica, the difference being particularly marked for Ca and Mg. P. radiata contained substantially higher concentrations of the metals Zn, Mn and Al than C. lusitanica, whereas the latter contained higher B concentrations. Possible reasons for differences between species in foliar nutrient concentrations are discussed.     

Effects of elevated nitrate and aluminum on the growth and nutrition of red spruce (Picea rubens) seedlings

Acidic deposition in high-elevation forests in the Appalachian Mountains of the eastern United States has been implicated in the decline of red spruce (Picea rubens Sarg.). Elevated soil acidity may increase soil Al availability and toxicity to roots. Enhanced soil solution NO(3) (-) concentrations, resulting from precipitation inputs and enhanced soil organic matter mineralization, may exacerbate Al toxicity by increasing root Al uptake. We exposed red spruce seedlings to 350, 500, 800 or 1400 micro M NO(3) (-) and 0 or 200 micro M Al in a factorial design in sand-nutrient solution culture to test if increased NO(3) (-) concentrations enhance Al uptake and toxicity. In addition to significant reductions in seedling growth parameters resulting from Al exposure, we found significant interactions between NO(3) (-) and Al for seedling height growth rate, needle weight, shoot weight and root weight. Differences in these parameters between Al treatments became more pronounced as solution NO(3) (-) concentration increased and reflected an Al-mediated inhibition of seedling response to increasing NO(3) (-) concentration. Solution NO(3) (-) concentrations above 500 micro M induced root nitrate reductase (NR) activity, whereas shoot NR activity increased in response to NO(3) (-) up to 500 micro M and declined above that concentration. In contrast, exposure to Al depressed NR activity of roots but tended to stimulate needle NR activity. Foliar N concentrations increased in seedlings grown in cultures containing between 350 and 500 micro M NO(3) (-), with no change above 500 micro M. Increasing concentrations of NO(3) (-) depressed foliar P concentrations, with reductions being greatest in seedlings exposed to 1400 micro M NO(3) (-). Exposure to Al increased foliar Ca, K and Al concentrations, decreased foliar P concentrations, and inhibited increases in foliar Mg concentration in response to increasing NO(3) (-). The consistent interactions between NO(3) (-) and Al for growth, root NR activity and foliar Mg concentration were the result of an inhibition of seedling response to NO(3) (-) mediated by Al in solution, rather than enhanced Al toxicity resulting from growth in the presence of elevated NO(3) (-) concentrations.     

Foliar nutrient composition in bareroot Pinus sylvestris nursery crops

We evaluated how the variability of nutrient concentrations affects their suitability as quality attributes. Foliar elements of four batches of three-year-old bareroot Scots pine seedlings, 150 seedlings from each batch, were analyzed. The variability in the concentrations of N, P, and K was small (coefficient of variation in a batch was 11–12), that of Ca and Zn intermediate (CV 19–21), and that of Mg, Mn, Fe, Al, Cu and Na was large (CV 26–35), being close to the variability in morphological attributes. Micronutrients varied more than macronutrients and a larger sample would be needed to estimate these concentrations with the same precision. Correlations between seedling size and nutrient concentrations in needles were small. Positive and negative correlations were found among nutrient concentrations, but not large enough to allow us to predict concentrations with each other. For concentrations of some nutrients the variance within batches was smaller than was found for morphological attributes in both relative and absolute terms, which provides support for batch culling.     

大花序桉幼苗营养吸收及分配规律研究

对大花序桉幼苗生物量及营养元素进行测定,结果表明,大花序桉苗木平均单株生物量大小排序为茎>根>叶。幼苗主要营养含量和吸收积累规律为K>N>P>Fe>Mn>Zn>B>Ca>Cu>Mg。大量元素N、P、K贮量最高的部位均为茎,根部和叶中含量相近。大花序桉苗期叶片对主要营养元素N、P和K的吸收分配间存在相互促进的关系;Mg与Ca,Cu与P、K,Zn与N、K、Cu、Fe,Fe与N呈现互相促进关系;Mn与N、Fe、K、Cu,B与P、Cu为相互抑制关系。据此,大花序桉苗期以氮、磷、钾肥为主,其中要求养分K₂O>N>P₂O₅,并有针对性地添加铁肥、锌肥和铜肥。     

Effects of complex stress of chronic acid fog exposure with rhizosphere aluminum on nutrient status in Abies firma seedlings

We exposed momi fir (Abies firma Sieb. et Zucc.) seedlings to simulated acid fog of pH 3.0 for about 3 years (from July 1999). During the last year of acid fog exposure, half of the seedlings were subjected to rhizosphere Al stress (complex stress) and the nutrient status of seedlings was determined. Chronic acid fog exposure decreased Fe and Zn concentrations in current-year and 1-year needles, and Al in 1-year needles, but had little effect on major element concentrations. Aluminum treatment had a broad impact on nutrient status in fine roots and needles. In fine roots, increases in Al and Cu concentrations and decreases in B, Mn, and Zn concentrations were observed. In 1-year needles, Al treatment increased Al, B, and Mn concentrations and decreased Cu concentration. The complex effect of acid fog with aluminum on nutrient status was relatively slight. These results show that changes in nutrient status under chronic acid fog exposure and/or Al stress are induced before the decline of photosynthesis in momi fir seedlings, mainly due to Al stress.     

Effects of chronic acid fog exposure with soil acidification or nitrogen loading on nutrient status in Abies firma seedlings

To clarify the nutrient status in momi fir (Abies firma Sieb. et Zucc.) seedlings under complex stress conditions of acid fog with soil acidification or nitrogen loading, we exposed seedling shoots to simulated acid fog (pH 3) and simultaneously loaded the soil with acid or excess nitrogen for 17 months. Proton and nitrogen loading reduced K concentration in soil, but these treatments had little effect on the nutrient status of fine roots in momi fir seedlings. Acid fog exposure resulted in reduced concentrations of Mg, Al, and Fe in current-year needles and Mg, Al, Cu, Fe, and Zn in 1-year-old needles. The complex effects of acid fog exposure with proton or nitrogen loading on nutrient status were relatively slight. However, elements such as Al and Cu notably reflect the effects of proton loading. These results show that the nutrient status of trace elements, rather than major elements, in needles of momi fir seedlings is sensitive to exposure to acid fog or proton loading.     

Organic Amendment,Fertilizer, and Lime Effects on Bareroot Jeffrey Pine Outplanted on a Sierra Nevada Surface Mine

Abstract

Composted organic matter, controlled release fertilizer, and dolomitic lime soil amendments were evaluated for their effectiveness in facilitating the reforestation of an acidic, semiarid Sierra Nevada surface mine with bareroot Jeffrey pine (Pinus jeffreyi Grev. & Balf.). Single application rates were used for the organic matter (2.0 L) and lime (28 g) amendments while low and high rates (8 g and 16 g, respectively) of the chemical fertilizer, High N 22-4-6 + Minors, were employed. All amendments were administered at outplanting using the minisite application method. The organic and lime amendments suppressed seedling survival, more so with the former than the latter, but this result did not extend to the High N fertilizer. Growth was also suppressed by the organic and lime amendments, although there was some evidence that these responses were waning late in the study, while High N was exceedingly stimulatory, especially with the high application rate. Annual assessment of seedling nutrition during this three-year study revealed numerous High N and organic matter treatment influences. Among them, increases in foliar N, P, and K and reductions in Fe, Mn, Zn, and Al were prominent within the High N treatments, but particularly so at the high application rate. For the organic matter treatment, the above increases and decreases in elemental concentrations were again noted, although the nutritional responses to this treatment were generally more subdued and somewhat ephemeral in comparison with those to High N. Calculation of base cation/metallic element molar ratios revealed that the organic matter treatment had the highest Ca/Al, Ca/Mn, Mg/Al, Mg/Mn, K/Al, K/Mn, and K/Cu during the initial season, but exerted little influence on these ratios thereafter. The High N treatments produced the highest Ca/Cu and Mg/Cu in the first season, the highest Ca/Al, Mg/Al, K/Al, and K/Mn in the second, and the highest Ca/Al, Mg/Al, Mg/Mn, K/Al, and K/Mn in the third season, with the high application rate especially prominent. Seedling nutrition was, at best, minimally affected by liming, with perhaps its most notable influence that of counteracting the positive responses indicated above with regard to the organic amendment effects on molar ratios. Overall, these results indicate that in the reforestation of difficult sites such as eastern Sierra Nevada surface mines, the favorable responses to controlled release fertilization can not be duplicated using composted organic amendments, and furthermore, dolomitic lime applications by the method used here are likely to prove exceedingly detrimental to seedling establishment.     

Effects of serpentine soil factors on Virginia pine (Pinus virginiana) seedlings

Effects of simulated serpentine soil conditions (elevated Mg:Ca ratio and Ni concentration) on seedlings from populations of Virginia pine (Pinus virginiana Mill.) from serpentine and non-serpentine sites were evaluated in sand culture. We determined (1) how seedlings are affected by elevated Mg:Ca ratio and Ni concentrations, (2) if there are interactive effects between Mg:Ca ratio and Ni concentrations on seedling growth, needle pigment concentrations, and nutrition, and (3) if Virginia pine populations from serpentine areas are edaphic ecotypes. A Mg:Ca ratio of 5 and 50 microM Ni both reduced seedling growth compared with control seedlings grown in the presence of the standard Mg:Ca ratio of 0.5 and no Ni. Interactive effects between Mg:Ca ratio and Ni concentrations were highly significant for growth, foliar pigments, and needle and root elemental concentrations. Nickel-mediated reductions in growth and foliar pigment concentrations were less at the serpentine Mg:Ca ratio of 5 than at the standard (non-serpentine) Mg:Ca ratio of 0.5. Foliar N was reduced by Ni concentrations as low as 10 microM, and foliar and root K, Ca and P concentrations were significantly reduced by Ni concentrations above 25 microM, with greater reductions at a Mg:Ca ratio of 0.5 than at a Mg:Ca ratio of 5. There were no population x serpentine soil factor interactions for seedling growth, foliar pigment concentrations, or nutrition, suggesting that seedlings from trees growing on serpentine soils are not edaphic ecotypes. We conclude that serpentine conditions present at the site of seed collection have not resulted in the selection of edaphic ecotypes of Virginia pine with respect to Mg:Ca ratio and Ni concentration.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.     

Acid mist and soil Ca and Al alter the mineral nutrition and physiology of red spruce

We examined the effects and potential interactions of acid mist and soil solution Ca and Al treatments on foliar cation concentrations, membrane-associated Ca (mCa), ion leaching, growth, carbon exchange, and cold tolerance of red spruce (Picea rubens Sarg.) saplings. Soil solution Ca additions increased foliar Ca and Zn concentrations, and increased rates of respiration early in the growing season (July). Soil Al treatment had a broad impact, reducing foliar concentrations of Ca, Mg, Mn, P and Zn, and resulting in smaller stem diameters, sapling heights and shoot lengths compared with soil treatments with no added Al. Aluminum treatment also reduced respiration when shoots were elongating in July and decreased net photosynthesis at the end of the growing season (September). Three lines of evidence suggest that Al-induced alterations in growth and physiology were independent of foliar Ca status: (1) Ca concentrations in foliage of Al-treated saplings were within the range of sufficiency established for red spruce; (2) mCa concentrations were unaffected by Al treatment; and (3) no Al x Ca interactions were detected. Acid mist treatment increased foliar Fe and K concentrations and increased leaching of Ca, Mg, Mn, Zn, Fe, and Al from foliage. Leaching losses of Ca were more than twice those of the element with the next highest amount of leaching (Zn), and probably led to the reductions in mCa concentration and membrane stability of acid-treated saplings. Acidic mist resulted in enhanced shoot growth, and consistent reductions in foliar cold tolerance in the fall and winter. Of the few significant interactions among treatments, most involved the influence of mist pH and Al treatment on foliar nutrition. In general, reductions in cation concentration associated with Al addition were greater for pH 5.0-treated saplings than for pH 3.0-treated saplings. We propose that H(+)-induced leaching of mCa from mesophyll cells is the mechanism underlying acid-induced reductions in foliar cold tolerance of red spruce.     

Urea fertilization and foliar nutrient composition of western hemlock (Tsuga heterophylla (Raf.) Sarc.)

Foliar mineral contents of hemlock stands in coastal and Cascade Mountain zones of the Pacific Northwest were studied to determine the effect of urea fertilization on foliar nutrients. The results demonstrate that mineral concentrations vary among foliage age classes, stands, and zones. Concentrations of N, P, K, Ca, Mg, Al, Mn, Fe, Cu, Zn and B were significantly higher in the Cascade Mountain stands than in the coastal stands. Urea fertilization significantly increased the N content in all age classes of foliage but reduced the concentrations of P, Ca, Mg, Mn, Fe, Al and B, especially in coastal stands. The poor response of coastal stands to urea fertilization, therefore, is probably due to further reduction in already low foliar levels of essential nutrients other than N.     

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.     

Growth and Nutritional Responses of Juvenile Jeffrey Pine to Post-Planting Fertilization

Abstract

Broadcast fertilization of a three-year-old Jeffrey pine (Pinus jeffreyi Grev. & Balf.) plantation growing on an eroded Sierra Nevada site was examined for its capacity to enhance seedling growth. The amendments chosen for the study included controlled release and conventional soluble formulations that contained a variety of N forms, but all featured high N content along with other nutrients. Four application rates were used, and all amendments were distributed around the base of the seedlings as top dressing. Four seasons after treatment application, fertilized seedlings were 129% larger in height, 93% larger in diameter, and 865% greater in shoot volume than unfertilized seedlings overall. Seedling sizes generally increased with application rate, and by a small margin, the controlled release fertilizer outperformed the other formulations. Relative growth analysis suggested that stimulation by the amendments would continue beyond four seasons. Foliar analysis conducted early and again late during the study revealed that N and P concentrations were consistently raised by fertilization, which likely accounts for much of the accompanying growth gains, while Ca, Fe, Mn, B, and Al concentrations were consistently reduced. Either early or late, foliar K was also raised while Mg, S, Zn, and Cu were reduced. Collectively, these results indicate that fertilization with a variety of high N formulations can increase the availability and uptake of critical nutrients sufficiently to impart in Jeffrey pine on low quality sites a substantial growth advantage.     

Distribution of elements along the length of Scots pine needles in a heavily polluted and a control environment

Pollution often causes visible symptoms of foliar injury. The injury is sometimes associated with an increase in the accessibility of toxic elements to plants as a result of acidification of the soil. We investigated the distribution of elements (N, P, K, Ca, Mg, Mn, S, Fe, B, Cu, Zn, Al, F, Pb, Cd, Cr, Ni and Co) in healthy current-year needles of Scots pine (Pinus sylvestris L.) growing at an unpolluted control site and at a site polluted mainly by SO(2), HF and Al(3+) from a fertilizer factory established in 1917. Needles from both sites were sampled before the appearance of visible injury and cut into five sections of equal length (tip, base and three middle parts). The mean concentrations of major nutrients were 20-30% lower in needles at the polluted site than in needles at the control site, whereas the concentrations of aluminum and fluorine were higher in needles at the polluted site. An increase in concentration from needle base to tip was detected for N, Fe, B, and Al at both sites and for Mn only at the polluted site. Fluoride accumulated in the tips of needles only at the polluted site, which could explain the necroses of needle tips at this site. The distribution of elements along the length of the needles was influenced by pollution, element mobility and the distal accumulation of toxic elements.