Effect of carbon dioxide, fertilization and irrigation on loblolly pine branch morphology

 Foliage and wood parameters of branches of 12-year-old loblolly pine (Pinus taeda L.) trees were characterized after 21 months of exposure to fertilizer, irrigation and elevated CO₂ treatments. Branches of loblolly pine trees were enclosed in plastic chambers and exposed to ambient, ambient +175 and ambient +350 umol mol^–1 CO₂ concentrations. Measurements of foliage and wood at the fascicle, flush and branch levels were made at the end of the 21 month study period. The +350 CO₂ treatment did not significantly increase fascicle radius or length but did increase the number of fascicles on the first flush. Fertilization significantly increased fascicle radius and length, while irrigation significantly increased number of fascicles and flush length of first flush. The +350 CO₂ treatment also significantly increased flush length of the first flush. Significant interaction of fertilization and irrigation with CO₂ was observed for fascicle length. Significant interactions of fertilization and irrigation were also observed for flush length, number of fascicles and fascicle length. Observed increases in fascicle radius, fascicle length, number of fascicles and flush length may have been responsible for the significantly higher flush leaf area observed for the all three treatments. Also, a combination of fertilization and irrigation increased leaf area by 82% compared to that in the control when averaged across CO₂ treatments. At the branch level +350 CO₂ treatment significantly increased shoot length but not the number of flushes on the branch. In general with the exception of bark density and total number of needle scales, neither fertilization nor irrigation had any significant effect on other branch level parameters. Results from this study indicate that with ‘global change’ an increase in CO₂ alone may increase leaf area via an increase in flush length and number of fascicles. Combining increases in CO₂ with fertilization and irrigation could greatly enhance leaf area which when coupled to observed increases in net photosynthesis as a result of elevated CO₂ could greatly increase productivity of loblolly pine trees. Received: 22 August 1996 / Accepted: 5 March 1997     

Testing the branch autonomy theory: a 13C/14C double-labelling experiment on differentially shaded branches

The impact of a heterogeneous within‐crown light environment on carbon allocation was investigated on young walnut trees trained on two branches: one left in full sunlight, the other shaded until leaf fall resulting in 67% reduction in photosynthetically active radiation. In September, the two branches were separately labelled with ¹⁴CO₂ and ¹³CO₂, respectively, so that the photosynthates from each branch could be traced independently at the same time. Although some carbon movements could be detected within 5 d in both directions (including from the shaded branch to the sun branch), between‐branch carbon movements were very limited: approximately 1% of the diurnal net assimilation of a branch. At this time of the year branch autonomy was nearly total, leading to increased relative respiratory losses and a moderate growth deficit in the shaded branch. The ratio of growth to reserve storage rate was only slightly affected, indicating that reserves acted not as a mere buffer for excess C but as an active sink for assimilates. In winter, branch autonomy was more questionable, as significant amounts of carbon were imported into both branches, possibly representing up to 10% of total branch reserves. Further within‐plant carbon transfers occurred in spring, which totally abolished plant autonomy, as new shoots sprouted on each branch received significantly more C mobilized from tree‐wide reserves than from local, mother‐branch located reserves. This allowed great flexibility of tree response to environment changes at the yearly time scale. As phloem is considered not functional in winter, it is suggested that xylem is involved as the pathway for carbohydrate movements at this time of the year. This is in agreement with other results regarding sugar exchanges between the xylem vessels and the neighbouring reserve parenchyma tissues.     

Patterns of branch permeability with crown depth among loblolly pine families differing in growth rate and crown size

Patterns in branch permeability with crown depth and permeability at the top of the main stem were analyzed for loblolly pine (Pinus taeda L.) trees from families selected on the basis of growth rate (fast, slow) and crown size (large, small). Analysis of variance with levels of crown size nested within levels of growth rate was used to test for differences in main stem permeability. Permeability at the top of the stem averaged 2.0×10^–12 m². There were no significant differences in permeability between families selected for fast and slow growth, but permeability was significantly lower for families selected for large crowns than for families selected for small crowns. Branch permeability averaged 0.74×10^–12 m² and decreased significantly with crown depth. Large-crown families had higher overall branch permeability than small-crown families. Average permeability in branches did not differ significantly between fast- and slow-growing families. Large crown-families had significantly larger current leaf area: total leaf area ratios in the lower two-thirds of the crown, and a weak but significant association was found between permeability and current leaf area: total leaf area ratios for a given relative crown depth. Our results suggest that ecotypic and adaptive processes simultaneously affect the overall patterns of stem and branch permeability in loblolly pine families.     

Differential shading of branches or whole trees: survival, growth, and reproduction

This study compared the effects of shading individual branches or whole trees on the survival, growth and reproduction of branches of mountain birch [Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti]. When a single branch was shaded its performance decreased drastically, while individual branches performed relatively well if the whole tree was shaded. For instance, after two summers 90% of the individually shaded branches were dead, while none of the study branches died following shading of entire trees. These results indicate that the fate of a branch depended more on shading-induced asymmetric competition among branches than on the shading itself. Furthermore, the increased growth of non-shaded branches in the trees where only two branches were shaded suggests that resources were preferentially allocated to branches in more favorable positions. Branch autonomy may promote environmental screening and growth towards favorable directions, optimizing the use of available light resources. In addition, branch autonomy may buffer the genet against environmental hazards, thus increasing the fitness of the genet at the cost of an increased mortality for individual modules.     

The effects of annual fertilization and complete competition control on fascicle morphology of different aged loblolly pine stands

The goal of this study was to determine how increased nutrient availability affects foliage morphology of loblolly pine (Pinus taeda L.) without introducing the confounding influence of light availability. Morphology of fascicles from the terminal leader (radius, length, specific needle area, density, and needles per fascicle) and terminal leader traits (length, foliated length, total leaf area, and total fascicles) were measured for different aged (ranged from 5-year-old to 12-year-old) loblolly pine stands growing on the Piedmont and lower Coastal Plain of Georgia, USA. A factorial combination of annual fertilization and complete interspecific competition control was applied as stand level treatments. Competition control and stand age generally did not affect fascicle morphology. In contrast, annual fertilization significantly increased fascicle length (5%), needles per fascicle (4%), and total leaf area of the terminal (18%), while decreasing specific needle area (4%). Fertilization also increased terminal leader growth (5%) and total number of fascicles (6%). Therefore, loblolly pine foliage morphology does have plasticity to respond to increased nutrient availability. However, the magnitude of these morphological changes is small compared to changes in total canopy leaf area.     

Gas exchange and canopy structure of 9-year-old loblolly pine,pitch pine and pitch x loblolly hybrids

Summary Seasonal gas exchange and canopy structure were compared among 9-year-old loblolly pine (Pinus taeda L.), pitch pine (Pinus rigida Mill.), and pitch x loblolly hybrids (Pinus rigida x taeda) growing in an F2 plantation located in Critz, Va., USA. Leaf net photosynthesis, conductance, internal CO₂ concentration (c_i), water use efficiency (WUE; photosynthesis/conductance), dark respiration and the ratio of net photosynthesis/respiration did not vary among or within the three taxa. Significant differences in volume production, crown length, total crown leaf surface area and the silhouette area of shade shoots among the taxa were observed. The loblolly-South Carolina source had greater volume and crown surface area than the pitch pine, and the hybrid taxa were intermediate between the two. Although the silhouette area ratio of shade foliage varied among taxa, it was not related to volume. A strong relationship between total leaf surface area and volume was observed. Leaf conductance, c_i, WUE and leaf water potential were the physiological parameters significantly and positively correlated with volume. This study suggests that the amount of needle surface in the canopy is more important in early stand volume growth than the leaf carbon exchange rate and the degree of needle self-shading in the lower canopy.     

Growth response of branches of Picea sitchensis to four years exposure to elevated atmospheric carbon dioxide concentration

Branch bags were used to expose branches on mature Sitka spruce trees to either ambient [CO₂] (A) or elevated [CO₂] (E) for 4 yr. This paper reports the effects of this treatment on the growth, development and phenology of the branches, including shoot expansion, shoot numbers, needle dimensions, needle numbers and stomatal density. The effect of elevated [CO₂] on the relationship between leaf area and sapwood area was investigated. Exposure to elevated [CO₂] doubled photosynthetic rates in current-year shoots and, despite some down-regulation, 1-yr-old E shoots also had higher rates of photosynthesis than their A counterparts. Thus, the amount of assimilate fixed by E branches was substantially more than that fixed by A branches; however, this increase in the local production of assimilate did not lead to an increase in non-structural carbohydrate or stimulate growth or meristematic activity within the E branches. There was a very consistent relationship between leaf area and stem cross-sectional area that was not influenced by [CO₂]. However, unbagged branches had thicker stems than bagged branches, resulting in a slightly lower ratio of leaf area to cross-sectional area. The implications of the results for the modelling of growth and allocation and the potential utility of the branch bag technique are discussed.     

Distribution of assimilate during the flush cycle of growth in Theobroma cacao L.

The growth of the shoot and roots of seedling plants of cocoa (Theobroma cacao L.) under constant glasshouse conditions showed a rhythmic cycle, with the maximum growth stages of each alternating in a regular sequence. When the growth cycle of the shoot was upset by removing all new leaves immediately after unfolding, the roots showed a high constant growth rate during this period, suggesting that normally the rapidly expanding leaves exert an inhibitory influence on the roots. Conversely removal of portions of the root delayed the production of new leaves in the shoot. The level of soluble and starch carbohydrate in the mature leaves, roots and stem declined during the period of expansion of the flush leaves, but accumulated again at the end of the leaf expansion stage. It is likely that this reserve carbohydrate was remobilised and translocated to the flush leaves during their period of expansion. A large proportion of newly formed photoassimilate, as shown by the distribution of ¹⁴C radioactivity from different source leaves, was also translocated to the young leaves during expansion. The large sink created by these leaves may cause photoassimilate and reserve carbohydrate to be diverted from the roots, thereby inhibiting root growth during the stage of leaf expansion. It is suggested that the rhythmic leaf production at the apex may control the growth cycle of the roots.     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

Influence of krummholz mat microclimate on needle physiology and survival

Summary Microclimate and photosynthesis of krummholz mat growth forms of Picea engelmanii (Parry) and Abies lasiocarpa [Hook.] Nutt. were investigated to determine structural features which may aid survival in alpine environments. The structure of krummholz mats was described in terms of the vertical distribution of leaf area index and leaf area density, which exceeded 50 m^-1 (based on total leaf surface area) near the canopy surface and approached zero below 30 cm from the surface in both species. Photosynthetic photon flux density (PPFD, 0.4–0.7 m wavelengths) and wind decreased by an average of 6 and 50-fold, respectively, between 1 m above and 10 cm below mat surfaces in both species. Needle temperatures on a P. engelmannii krummholz mat during July averaged about 2°C above air temperature during the day, with a maximum overtemperature of greater than 20°C above T _air during one sunlit period. At night, needle temperatures averaged 3–4°C below T _air.Net photosynthesis in year-old P. engelmannii shoots reached a maximum at 15–20°C during July and August. Surface shoots were light saturated at near 1200 moles m^-2s^-1 PPFD, and had higher photosynthetic rates than subsurface, predominantly shaded shoots above 800 moles m^-2s^-1. Shade shoots had higher photosynthetic rates when PPFD was below 600 moles m^-2s^-1, and at 250 moles m^-2s^-1 shade shoots maintained about 50% of the net photosynthetic rate of sun shoots at light saturation. Shade shoots appeared capable of benefitting photosynthetically from elevated temperatures within krummholz mats despite relatively low light levels. Especially rapid photosynthesis may occur when canopy needles are illuminated by sunflecks and needle temperatures rise by 10° C or more.Snow cover appears crucial for the survival of needles during winter. Snow accumulated within krummholz needle canopies before the sub-canopy zone of unfoliated branches became filled. The concentrated needle growth in the krummholz canopy captured snow in early autumn without support from ground-level snowpack. Early snow cover in both species prevented cuticle abrasion and resulted in high winter needle water contents and viabilities for subsurface compared to surface needles which became abraded, severely dehydrated, and had high mortality between December and February, especially on windward sides of shoots.Extremely high concentrations of needles within krummholz mat canopies created an aerodynamic structure which elevated needle temperatures to more optimal photosynthetic levels in summer and resulted in more efficient snow accumulation in winter. These factors appear crucial for winter needle survival. Thus, krummholz mats appear to be an important adaptation in growth form which provides survival benefits in both summer and winter.     

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.     

Micropropagation of shortleaf,Virginia and loblolly x shortleaf pine hybrids via organogenesis

Protocols were developed for the micropropagation of shortleaf pine (Pinus echinata Mill.), loblolly (P. taeda L.) x shortleaf pine hybrids, and Virginia pine (P. virginiana Mill.). For meristematic tissue induction, modified Gresshoff & Doy (GD) medium with a high concentration of benzyladenine (BA) and short pulse treatment was best for loblolly x shortleaf hybrids whereas a lower concentration of BA and longer pulse treatment was best for shortleaf and Virginia pines. Shoot growth rate for all species was generally slower on Schenk & Hildebrandt medium than on GD medium. Addition of activated charcoal improved shoot growth of shortleaf pine but not of Virginia pine or the loblolly x shortleaf hybrids. Separation of shoots was beneficial before placing in the advanced growth medium. Both GD and Litvay's media produced good advanced shoot growth, especially following the addition of 0.5% activated charcoal. Individual shoot heights of 2–3 cm and 8–12 weeks of age after separation from the cluster were best for rooting. Root induction declined rapidly thereafter. Modified GD medium with 0.5 mg 1^-1 -naphthaleneacetic acid plus 1.0 mg 1^-1 3-indolebutyric acid and 20 g 1^-1 sucrose was best for root induction for all species except shortleaf pine. Addition of activated charcoal produced better root systems. Too high a light intensity resulted in a lowered frequency of rooting. A large number of plantlets was produced.     

Effects of elevated CO2-grown loblolly pine needles on the growth, consumption, development, and pupal weight of red-headed pine sawfly larvae reared within open-topped chambers

Seedlings of loblolly pine, Pinus taeda , were grown in open-topped chambers under four levels of CO₂: two ambient and two elevated. Larvae of the red-headed pine sawfly, Neodiprion lecontei , were reared from early instar to pupation, primarily on branches within chambers. Larval growth and mortality were assessed and leaf phytochemistry samples of immature and mature leaves collected weekly. Mature leaves grown under elevated CO₂ had significant reductions in leaf nitrogen and increases in non-structural carbohydrate contents, resulting in foliage being a poorer food source for larvae, i.e. higher carbohydrate:nitrogen ratio. Nutritional constituents of immature needles were unaffected by seedling CO₂ treatment. Volatile mono- and sesquiterpenes were unrelated to plant CO₂ treatments for either leaf age class. Larval consumption of immature needles significantly increased on seedlings grown under CO₂ enrichment, while mature needle consumption was not different between the treatments. The average weight gain per larva significantly declined in late instar larvae consuming elevated CO₂-grown needles. In spite of this reduced growth, neither the days to pupation nor pupal weights were different among the CO₂ treatments. This study suggests that enriched CO₂-induced alterations in pine needle phytochemistry can affect red-headed pine sawfly performance. However, compensatory measures by larvae, such as choosing to consume more nutritious immature needles, apparently helps offset enriched CO₂-induced reductions in the leaf quality of mature needles.     

Uniform versus Asymmetric Shading Mediates Crown Recession in Conifers

In this study we explore the impact of asymmetrical vs. uniform crown shading on the mortality and growth of upper and lower branches within tree crowns, for two conifer species: shade intolerant lodgepole pine (Pinus contorta) and shade tolerant white spruce (Picea glauca). We also explore xylem hydraulics, foliar nutrition, and carbohydrate status as drivers for growth and expansion of the lower and upper branches in various types of shading. This study was conducted over a two-year period across 10 regenerating forest sites dominated by lodgepole pine and white spruce, in the lower foothills of Alberta, Canada. Trees were assigned to one of four shading treatments: (1), complete uniform shading of the entire tree, (2) light asymmetric shading where the lower 1/4–1/3 of the tree crown was shaded, (3) heavy asymmetric shading as in (2) except with greater light reduction and (4) control in which no artificial shading occurred and most of the entire crown was exposed to full light. Asymmetrical shading of only the lower crown had a larger negative impact on the bud expansion and growth than did uniform shading, and the effect was stronger in pine relative to spruce. In addition, lower branches in pine also had lower carbon reserves, and reduced xylem-area specific conductivity compared to spruce. For both species, but particularly the pine, the needles of lower branches tended to store less C than upper branches in the asymmetric shade, which could suggest a movement of reserves away from the lower branches. The implications of these findings correspond with the inherent shade tolerance and self-pruning behavior of these conifers and supports a carbon based mechanism for branch mortality – mediated by an asymmetry in light exposure of the crown.     

Branch transpiration of pine and spruce scaled to tree and canopy using needle biomass distributions

Branch water exchange and total tree water uptake were measured in a mixed Norway spruce and Scots pine stand in central Sweden during the 1995 and 1996 growing seasons. Branch transpiration was scaled to canopy level on the basis of a branch conductance model, using vertical needle-area distributions obtained by destructive sampling. Comparison with total tree water uptake scaled to canopy level showed agreement within 10%, for periods when the canopy was not affected by climatically induced stress. Comparison of scaled fluxes on individual trees showed that measurements of transpiration at branch level provide information on the direct response of transpiration to variations in weather, and furthermore that the time-lag between transpiration and tree water uptake was as much as 3 h. The vertical needle-area distribution of Scots pine was similar to that found by other authors. Needle-area distribution on Norway spruce, which has not been described before, showed that it has its largest needle area at the top of the crown. Specific needle area varied considerably both within trees and between trees. For spruce, mean specific needle area (±SD) varied from 2.4±0.5 mm² mg^–1 at the top of the crown to 7.1±1.9 mm² mg^–1 at the base. Corresponding figures for Scots pine were 3.4±2.0 and 9.1±2.1. Received: 5 March 1999 / Accepted: 17 March 2000     

Photosynthetic capacity,chloroplast pigments,and mineral content of the previous year's spruce needles with and without the new flush: analysis of the forest-decline phenomenon of needle bleaching

Summary Spruce (Picea abies) damage in the Fichtelgebirge (FRG) occurs as needle bleaching and a depression of CO₂ assimilation. Such injury may primarily result from the direct, above-ground effects of air pollution or indirect, below-ground changes in mineral uptake.Typically, the new flush of spruce needles is green and exhibits high photosynthetic capacity. Mies and Zöttl concluded that the older foliage is damaged when nutrients are withdrawn to supply the current year's needles. By removing the terminal buds of single branches in the spring, we produced an experimental set of the previous year's needles with greater mineral reserves than the control needles. During the course of the growing period, the performance of the experimental needles, which lacked competition from the new flush, was compared to that of the control needles of the same age-class on intact branches with the new flush.Throughout the experiment, chloroplast pigments of a healthy control tree were not affected by the elimination of the new flush. However, the chlorophyll and carotenoid content as well as the photosynthetic capacity of the previous year's needles on those branches of a heavily damaged tree where the new flush had been eliminated increased substantially. This increase was associated with an increase in minerals, which seemed to be deficient in the control needles with the new flush. Thus, in contrast to needles of the same age-class on intact branches with undisturbed new growth in the same atmospheric environment, the experimental needles escaped bleaching and a decrease in photosynthesis. It would seem that the bleaching and the loss in photosynthetic capacity typical of trees damaged by forest decline indirectly result from nutrient deficiencies through soil environment changes and/or root damage than directly from atmospheric pollutants.     

Seed source variation in 32P uptake in inus taeda L. seedlings: A possible regulation by efflux

Short-term ³²P uptake experiments were conducted with intact seedlings of loblolly pine (Pinus taeda L.) to examine possible seed source variation in net accumulation of ³²P in roots and shoots, and in rates of unidirectional influx. Seed source had a highly significant effect on biomass and P concentrations of shoots and roots. Seedlings from two seed sources representing fast-growing populations (a broadly-adapted and wet-site seed source) accumulated over 60% more total seedling P than smaller seedlings from a drought-hardy seed source, reflecting higher biomass and root P concentrations. Rates of unidirectional ³²P influx in seedlings from the drought-hardy seed source were more than twice the rates of the seedlings from the broadly-adapted seed source. However, after 24 h in labeled uptake solution, net accumulation of ³²P was similar, suggesting that rates of unidirectional efflux from roots of the drought-hardy seed source were also high. Although there were no significant differences in biomass and tissue P concentrations between the two fast-growing seed sources, rates of unidirectional influx in seedlings from the broadly-adapted seed source were 42% lower than rates in seedlings from the wet-site source. Yet, after 24 h in labeled uptake solution, net accumulation of ³²P in seedlings from the broadly-adapted seed source was 50% higher. Unidirectional efflux out of the root may regulate net uptake of P as much, if not more, than influx in loblolly pine seedlings-at least under high-P growth conditions. The results in this study do not support previous studies with herbaceous plants suggesting that fast-growing species typically exhibit higher rates of nutrient uptake than slow-growing species.     

Phloem transport in Picea abies (L.) Karst. in mid-winter

Summary Translocation of ¹⁴C assimilates was studied on four different transport systems of Picea abies branches after induced activation in January. ¹⁴CO₂ assimilation of terminal shoots for 48 h at 25° C resulted in phloem loading and basipetal transport of ¹⁴C photosynthate into the following, older shoot generations. ¹⁴C import was enhanced, when these older shoot generations were kept in the dark. Microautoradiographs of the labelled terminal shoots showed that ¹⁴C assimilates were exported from needles via sieve elements of the leaf traces and loaded into the latest increment of the axial secondary phloem. No ¹⁴C label appeared in the obliterated sieve cells or in the tracheids. In addition, ¹⁴C photosynthate accumulated densely in the chlorophyllous cells of the cortex and in cells of the resin ducts, indicating certain sink activity. In the darkened 2-year-old shoot, imported ¹⁴C photosynthate was concentrated in the functional secondary phloem, while some ¹⁴C label was unloaded into the latest xylem increment. When 6-year-old shoots were exposed to ¹⁴CO₂ for 48 h in the light, ¹⁴C assimilates accumulated in the phloem of the leaf trace and in the latest increment of the axial secondary phloem. However, a substantial amount of radioactivity was unloaded into ray cells and phloem parenchyma cells. Thus, the presence of functioning phloem in needles and twigs of P. abies during winter allows long-distance translocation and radial distribution of assimilates according to existing source-sink relations.     

Phenotypic analysis of first-year traits in a pseudo-backcross {(slash x loblolly) x slash} and the open-pollinated families of the pure-species progenitors

A single test, including one pseudo-backcross (Pinus elliottii x Pinus taeda) x P. elliottii and open-pollinated families of the pure species progenitors, was established in North Central Florida in December 2007 to study the transfer of the fast-growing characteristics from a P. taeda L. (loblolly pine) parent into the P. elliottii Engelm. (slash pine) background. Several traits were measured in the first growing season: height growth, phenology, tip moth incidence, stem traits, crown architectural and needle traits. Heterosis was evaluated for each trait using analyses of variance by fitting a linear mixed model. All traits were significantly (p value < 0.05) different among families while the significance for heterosis varied by trait. Positive heterosis was found for average rate of shoot elongation (ASRE), total growth (TG), total height and number of needles per fascicle while the opposite was true for base diameter, top diameter, fascicle length, fascicle diameter, crown projected area and phenological traits (cessation, duration and day to reach 50% of the height). Average performance (i.e., no heterosis) was found for initiation of growth, number of branches, number of nodes, tip moth incidence, sheath length and specific leaf area. The analyses indicated that introgression of loblolly pine alleles into slash pine was effective and novel trait combinations were achieved. The pseudo-backcross had larger variation in early height growth than the slash pine families and was taller than all open-pollinated families at the end of the first season. Tip moth incidence was much lower than the loblolly pine family.     

Photosynthetic performance,chloroplast pigments,and mineral content of various needle age classes of spruce (Picea abies) with and without the new flush: an experimental approach for analysing forest decline phenomena

Summary Damage in the older needles of Norway spruce [Picea abies (L.) Karst.] in the Fichtelgebirge (NE Bavaria, FRG) appears to result primarily from nutrient imbalances rather than from direct effects of air pollutants on the mesophyll of the needles. Support for this conclusion was obtained by altering the nutrition of older needles through the removal of terminal buds on several branches from a damaged and an undamaged spruce tree in spring. Various photosynthetic parameters, as well as the chloroplast pigment and nutrient concentrations, of 1- to 3-year-old needles on manipulated branches were compared with those of branches on which the new flush was allowed to develop during the course of the growing period. Removal of terminal buds affected only the 1-year-old needles. Elimination of the new flush resulted in a higher Ca and Mn content of the needles of the undamaged tree. This treatment also resulted in an increase of the photosynthetic capacity (under saturating light and CO₂ conditions), carboxylation and light use efficiency, as well as net photosynthesis under natural conditions of the 1-year-old needles on the yellow chlorotic tree. This was accompanied by higher chlorophyll concentrations and an increase in Mg, Ca, Mn, and Zn content, and no visible signs of chlorosis developed in the experiment. By contrast, the needles of twigs in which the new flush was allowed to develop exhibited reductions in mineral content in the middle of the year. This was especially true for the elements Mg and Ca, and was accompanied by needle chlorosis and a depression of the capacity of photosynthesis. Thus it appears that there is a close relationship between the development of needle damage and nutrient imbalances in spruce. The retranslocation of elements from the 1-year-old needles to the new flush seems to play a major role in the development of needle bleaching. This approach thus supports the hypothesis described above and confirms a preliminary test with a similar experimental design, which had been conducted earlier.