Mobilization and Incorporation of Photoassimilated C by Growing Vegetative and Reproductive Tissues of Adult Pinus resinosa Ait. Trees

Variations in translocation and incorporation of ¹⁴C-labeled photosynthate in actively growing vegetative and reproductive tissues of 20-year-old Pinus resinosa trees with time were investigated. Five hours after exposure of 1-year-old needles on second order branches to ¹⁴CO₂, 90% of the recoverable activity was still in the needles. The 1-year-old bark showed a small amount of activity, but terminal sinks had accumulated little ¹⁴C. After 24 hours, activity in the 1-year-old needles decreased to 59% while actively growing terminal and lateral shoots and 2nd year cones had accumulated considerable tracer. Whereas activity in the 1-year-old bark increased, recovery from the 1-year-old wood was only 1.8%. After 1 week, relative activity in the 1-year-old needles had declined to 27%. Recovery of ¹⁴C from various sinks was high and in the following order: 2nd year cones > terminal needles > lateral needles > terminal internode > lateral internodes > 1-year-old wood. Ninety per cent of the ¹⁴C recovered after 5 hours was in the ethanol-soluble form, but by the end of 1 week only half was in the soluble fraction. Incorporation of tracer into insoluble compounds was high in expanding internodes and in 1-year-old xylem, and low in 2nd year cones. Total activity in the terminal internode generally declined toward the distal end while specific radioactivity increased. Only 5% of the ¹⁴C recovered from cones was localized in developing seeds, but specific radioactivity of seeds was higher than in the cone scales and axis.     

Carbon autonomy of reproductive shoots of Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var.<Emphasis Type="Italic"> sibirica</Emphasis>)

Carbon autonomy of current-year shoots in flowering, and of current-year shoots plus 1-year-old shoots (1-year-old shoot system) in fruiting of Siberian alder (Alnus hirsuta var. sibirica) was investigated using a stable isotope of carbon, ¹³C. The current-year shoot and 1-year-old shoot systems were fed ¹³CO₂ and the atom% excess of ¹³C in flowers and fruits was determined. The majority of photosynthate allocated to flower buds was originally assimilated in the leaves of the flowering current-year shoots. Of all the current-year shoots on fruiting 1-year-old shoots, only those nearest to the fruits allocated the assimilated photosynthate to fruit maturation. These results indicate that the current-year shoots and 1-year-old shoot systems are carbon-autonomous units for producing flowers and maturing fruits, respectively.     

Control of Adventitious Root Production and Hypocotyl Hypertrophy of Sunflower (Helianthus annuus) in Response to Flooding

Needles of 20-year-old Scots pine trees (Pinus silvestris L.) were permitted to photoassimilate ¹⁴CO₂ for 1 h on different dates during the growing season. The loss of radioactivity from current, 1-year-old, and 2-year-old needles was followed, and the translocation of photoassimilated ¹⁴CO₂ from older needle age-classes to the elongating new needles studied. The effects of good mineral and water supply on translocation were also considered. In the spring, 1-year-old and 2-year-old needles accumulated ¹⁴C. These reserves, together with current photosynthate, were utilized when the trees started growing. The 1-year-old needles exported ¹⁴C to the current needles during the first weeks of elongation of the later, while no such translocation occurred from the 2-year-old needles. Removal of the 1-year-old needles resulted in translocation of assimilates from the 2-year-old needles to the current needles. The general pattern of translocation observed in the control trees was not changed when the trees were fertilized and irrigated. The new needles started to export assimilates in the middle of July when the photosynthetic rate per needle was comparable with that of the older age-classes. This occurred about 4 weeks after positive net photosynthesis was first measured for the current shoot. The current needles of trees with good nutrient and water supply seemed to become self-sufficient in photoassimilates earlier than the current needles of the control trees.     

Seasonal changes in the utilization and turnover of assimilation products in 8-year-old Scots pine (Pinus sylvestris L.) trees

Summary The present study aimed at a physiological understanding of the seasonal changes of the carbohydrate patterns and levels in the various tissues of 8-year-old Scots pine (Pinus sylvestris L.) trees growing under ambient climatic conditions in the botanical garden at Bayreuth. The photosynthates of selected twig sections were labelled by ¹⁴CO₂ fixation and after chase periods of 1 h up to 8 months, the distribution of radiocarbon in the whole trees was determined and the labelling of identified carbohydrates was compared with the levels of these compounds in the individual tissues. Bud break and sprouting in spring is exclusively supplied by the recent photosynthates of the previous year's needles. During summer assimilates of the old needles were utilized for secondary growth of the axial system while growth of the recent-year's shoots was supported by their own photosynthesis. In autumn, soluble carbohydrates were produced instead of starch, a major part of which in addition to recent photosynthates was utilized for root growth during the cold season. Another part of the autumnal storage material was incorporated into the cell walls of the latest xylem and phloem elements still in winter. A pronounced starch-oligosaccharide interconversion upon frost hardening, and its reversal in spring as has been described for deciduous trees, could not be observed. This was due to maintenance of photosynthetic capability even in the cold season and the replacement of consumed storage material especially in late winter and early spring by new photosynthates.     

Effect of defoliation on tracheid production and the level of indole-3-acetic acid in Abies balsamea shoots

To simulate feeding by the spruce budworm ( Choristoneura fumiferana Clem.), the apical current-year shoots on 1-year-old branches in the uppermost whorl of 6-year-old balsam fir [ Abies balsamea (L.) Mill.] trees were either removed completely by debudding before the start of the growing season or defoliated 0, 50, 90 or 100% shortly after budbreak. Debudded branches were treated at the apical end with 0, 0.1 or 1.0 mg of indole-3-acetic acid (IAA) (g lanolin)⁻¹. Ninety % of the 1-year-old needles were also removed from some of the experimental branches. After ca 4 weeks of growth, the radial width of new xylem and the level of IAA were determined in the 1-year-old internode. The IAA content was measured by radioimmunoassay.
The removal or defoliation of current-year shoots inhibited tracheid production and decreased the IAA level. Exogenous IAA stimulated tracheid production and increased the IAA level in debudded branches. Current-year shoot defoliation also inhibited current-year shoot elongation. The inhibitory effect of current-year needle removal on all parameters generally increased with increasing intensity of defoliation. The removal of 1-year-old needles did not affect the IAA level or current-year shoot elongation, nor did it influence tracheid production in branches with current-year shoots. However, removal of 1-year-old needles inhibited tracheid production in debudded branches supplied with exogenous IAA. The results indicate that (1) IAA is involved in the control of tracheid production in the 1-year-old internode, (2) IAA is supplied primarily by current-year shoots, and (3) defoliation by the spruce budworm inhibits tracheid production partly by decreasing the supply of IAA.     

Seasonal Changes in the Distribution of Photo-assimilated C in Young Pine Plants

Pinus strobus L. plants in their third year of growth were permitted to photoassimilate ¹⁴CO₂ for about 1 hour at monthly intervals between April and October, and the subsequent distribution of ¹⁴C in these plants was determined 8 hours, 1 month, 2 months or 4 months after photo-assimilation. In this way, the fate of ¹⁴CO₂ photo-assimilated during different months of the growing season was observed.

In the spring, old needles played a significant role in photo-assimilating ¹⁴CO₂ and exporting current photosynthate to the developing new shoots and roots. By July, the new shoot had replaced the old shoot both as the primary photo-assimilating part of the plant and as an exporter, particularly to the root.

The root received current photosynthate from the shoot throughout the entire growing season, although plant analysis only 8 hours after photo-assimilation did not always reveal this. Translocation of recent photosynthate from shoot to root was particularly high in August, September, and October.

The amounts of photo-assimilated ¹⁴C lost from the plants over a 4 month interval, principally through respiration and photorespiration, were about one-half of that absorbed during photo-assimilation, with the greatest loss occurring within the first month.

     

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.     

LEAF TURNOVER RATES OF ADENOSTOMA FASCICULATUM (ROSACEAE)

The age structure of the foliage of a 26-yr-old stand of Adenostoma fasciculatum H. & A. (chamise) was analyzed. The mean number of standing leaves and the yearly increase in leaf scars on the leaf-producing short shoots allowed an estimate of annual leaf production. The average chamise leaf persists for two seasons. Short shoots produce 4–6 leaves per yr; however after 4–5 yr their productivity declines. About 72% of the standing leaves were produced during the current and 28% during the foregoing season. Nearly one-half of all the leaves produced was found on current-year short shoots (i.e., on long shoots that had developed during the spring of the same year). Thus, earlier estimates of leaf production in chamise based only on current-year long shoot growth were too low.     

Effects of previous defoliation on pine looper larval performance

1 During outbreaks of the pine looper, Bupalus piniarius, its host, Pinus sylvestris, is severely defoliated. The larvae of this geometrid normally feed on mature needles. However, because trees are totally defoliated during outbreaks, the next generation is forced to feed on current-year needles. 2 Bupalus piniarius larvae feeding on previously defoliated trees may show lower performance either because of induced resistance or because larvae have to feed on needles not normally fed upon (current instead of mature). 3 These hypotheses were tested in an experiment where larvae were reared on (i) shoots naturally defoliated the previous year, and thus, bearing only current-year needles, (ii) non-defoliated shoots where larvae had access only to current-year needles, and (iii) control shoots with access to both current and mature needles. 4 There was no support for the induction hypothesis. Survival was lower on naturally defoliated shoots than on control shoots (81.3 vs. 90.9%), but survival was lower also on non-defoliated shoots where larvae had access only to current-year needles (78.8%). Data on larval feeding distribution showed a strong preference for mature needles. 5 Needle nitrogen concentration of current-year needles was 38% higher on defoliated trees than on non-defoliated trees. 6 It is concluded that defoliation affected larval performance primarily through the removal of the preferred type of needles and not because of an induced resistance. Effects of increased concentrations of allelochemicals in defoliated shoots, if present, were probably cancelled out by increased nitrogen concentrations.     

Carbohydrate Accumulation in Siberian Spruce Needles of Various Ages

The content of mono- and oligosaccharides, hemicelluloses, and water-soluble polysaccharides was studied in Siberian spruce (Picea obovata Ledeb.) needles of various ages. Accumulation of carbohydrates during the growth period and the ratio of their individual fractions were determined by the proportions between the growth rate and assimilate export in developing needles. Throughout the growth period, the content of monosaccharides in needles of all ages was much higher than that of oligosaccharides. Unlike the period of active growth, the dormancy period was characterized by a wider diversity of oligosaccharides and their rather high content. Hemicelluloses that accumulated during the previous growth season and water-soluble polysaccharides of 2- and 3-year-old needles were involved into metabolism at the time of active growth of current-year needles. Analysis of carbohydrate accumulation in 1- to 9-year-old needles showed that the monosaccharide concentration was higher in needles of the first two years. The content of hemicelluloses A and B and water-soluble polysaccharides was higher in older needles. During the growth period, the assimilates of 2- and 3-year-old needles were partly used for the formation of young needles. The increase in relative content of polysaccharides during needle senescence reflects the involvement of photosynthetic products into the secondary pool of carbon metabolites.     

Quantitative analysis of carbon balance in the reproductive organs and leaves of <Emphasis Type="Italic">Cinnamomum camphora</Emphasis> (L.) Presl

We investigated seasonal changes in dry mass and CO₂ exchange rate in fruit and leaves of the evergreen tree Cinnamomum camphora with the aim of quantitatively determining the translocation balance between the two organs. The fruit dry mass growth peaked in both August and October: the first increase was due to fruit pulp development and the second to seed development. Fruit respiration also increased with the rapid increase in fruit dry mass. Therefore, the carbohydrates required for fruit development showed two peaks during the reproductive period. Fruit photosynthesis was relatively high in early August, when fruit potentially re-fixed 75% of respired CO₂, indicating that fruit photosynthesis contributed 15–35% of the carbon requirement for fruit respiration. Current-year leaves completed their growth in June when fruit growth began. Current-year leaves translocated carbohydrates at a rate of approximately 10–25 mg dry weight (dw) leaf⁻¹ day⁻¹ into other organs throughout the entire fruit growth period. This rate of translocation from current-year leaves was much higher than the amount of carbohydrate required for reproduction (ca. 3 mg dw fruit⁻¹ day⁻¹). Given the carbon balance between fruit and current-year leaves, carbohydrates for reproduction were produced within the current-year fruit-bearing shoots. C. camphora would be adaptive for steadily supplying enough amount of carbohydrate to the fruits, as there was little competition for carbohydrates between the two organs. As assimilates by leaves are used for processes such as reproduction and the formation of new shoots, photosynthesis by reproductive organs is considered to be important to compensate for reproductive cost.     

Stem-growth response of Pinus sylvestris and Picea abies to nitrogen fertilization as related to needle nitrogen concentration

 Responses of stem-volume growth to N application were evaluated in relation to foliar N concentrations. Data from N-fertilization experiments in 28 Pinus sylvestris stands and 21 Picea abies stands were used. Relative stem-growth responses were negatively related to concentrations of N in current-year needles of unfertilized trees. There appeared to be a threshold value of 15–16 mg (g DM)^–1 N in current-year needles, above which N-application is unlikely to stimulate growth. However, relations were non-significant between N concentrations in current-year needles and the absolute stem-growth response [dm³ ha^–1 (5 years)^–1]. The indicated threshold values are discussed in relation to other variables reflecting the N richness of the environment.--> Received: 20 December 1996 / Accepted: 30 September 1997     

Compensatory photosynthesis as a response to partial debudding in ezo spruce, <Emphasis Type="Italic">Picea jezoensis</Emphasis> seedlings

The effects of partial debudding on photosynthesis, stomatal conductance and nitrogen contents of 1-year-old needles and newly grown needles were studied in Picea jezoensis (Sieb. et Zucc.). Seventy-five percent of the buds of P.jezoensis seedlings were removed soon after bud break. Gas exchange was measured three times for 1-year-old needles and twice for current-year needles. The photosynthetic rates of 1-year-old needles were greater in debudded seedlings than in control seedlings, and the difference increased as the growing season progressed. This greater photosynthetic rate in debudded seedlings was accompanied by greater stomatal conductance. However, neither the photosynthetic rates nor the stomatal conductance of current-year needles differed between debudded and control seedlings after the needles had fully expanded. Debudding also had no effect on mass-based nitrogen contents in either the 1-year-old or the current-year needles. Area-based nitrogen in the 1-year-old needles did not differ between debudded and control seedlings, but was greater in debudded seedlings than in control seedlings in current-year needles. These results suggest that the enhanced photosynthetic rate is more likely a result of an increased root/leaf ratio that reduces the stomatal limitation of photosynthetic rate than a result of altered sink-source relationships or increased leaf nitrogen content.     

Variation of specific leaf area and upscaling to leaf area index in mature Scots pine

Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm² g⁻¹) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m² m⁻²) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.     

Effects of Fertilization and Irrigation on the Seasonal Changes of Carbohydrate Reserves in Different Age-Classes of Needle on 20-Year-Old Scots Pine Trees (Pinus silvestris)

The effects of fertilization, irrigation or both on the seasonal changes of starch and soluble carbohydrates (glucose, fructose, myo-inositol, pinitol and sucrose) in needles of 20-year-old Scots pine trees (Pinus silvestris L.) were studied during three consecutive years. The starch content of the mature needles increased during spring and early summer to about 25% of dry weight. Neither fertilization nor irrigation affected the general pattern of starch accumulation during the spring. The starch reserves were mobilized when the shoot started to grow. Starch content decreased more rapidly in needles from fertilized than in those from unfertilized trees. The current needles from the control trees accumulated starch while they were still growing. The current needles of the fertilized trees did so to a lesser extent. The amount of starch was closely correlated to the air temperature and to the growth rate. Large amounts were found at low temperatures and low growth rates. The concentrations of soluble carbohydrates showed the well-known seasonal variation, with the highest value during the winter. The levels of sugars were nearly similar, irrespective of fertilization. An exception was sucrose, which was found in small quantities in needles from fertilized plots. Small amounts of sucrose were also found in growing current needles. The results are discussed in relation to growth limitation by assimilate availability and indicate that the ‘sink demand’ is the limiting factor.     

Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species

In trees, reproduction constitutes an important resource investment which may compete with growth for resources. However, detailed analyses on how growth and fruit production interact at the shoot level are scarce. Primary canopy growth depends on the development of current-year shoots and their secondary growth might also influence the number and size of fruits supported by them. We hypothesise that an enhanced thickening of current-year shoots is linked positively to acorn production in oaks. We analysed the effect of acorn production on shoot growth of two co-occurring Mediterranean oak species with contrasting leaf habit (Quercus ilex, Quercus faginea). Length and cross-sectional area of current-year shoots, apical bud mass, number of leaves and acorns, xylem and conductive area, number of vessels of acorn-bearing and non-bearing shoots were measured in summer and autumn. Nitrogen and carbohydrates analyses were also performed in stems and leaves of both shoot types. Stem cross-sectional area increased in acorn-bearing shoots when compared with non-bearing shoots for both species and such surplus secondary growth was observed since summer. In bearing shoots, the total transversal area occupied by vessels decreased significantly from basal to apical positions along the stem as did the xylem area and the number of vessels. Leaves of bearing shoots showed lower nitrogen concentration than those of non-bearing shoots. Carbohydrate concentrations did not differ in stems and leaves as a function of the presence of acorns. Such results suggest that carbohydrates may preferentially be allocated towards reproductive shoots, possibly through enhanced secondary growth, satisfying all their carbon demands for growth and reproduction. Our findings indicate that acorn production in the two studied oaks depends on shoot secondary growth.     

Involvement of Exported Photosynthetic Products in the CO2Exchange of the Skeletal Shoots of Pine

CO₂exchange in the leafy and skeletal parts of attached shoots of Pinus sylvestrisL. was measured with an infrared gas-analyzer in an open differential system during daylight hours. The ¹⁴CO₂assimilation rates in the leafy parts of shoots and ¹⁴CO₂evolution from current photosynthetic products in the lower skeletal part of shoots were measured in afternoons. Chlorophyll content was measured in the needles of the same shoot. The carbon of exported assimilates contributed only about 4% to CO₂exchange in the heterotrophic tree tissues. Only this component of CO₂evolution from the surface of the skeletal part of the tree was related to the losses of the net primary photosynthetic production (NPP) in the aboveground part of the pine stand during the current growth period.     

Seasonal Course of Photosynthesis, Respiration, and Distribution of C in Young Pinus resinosa Trees as Related to Wood Formation

Rates of net photosynthesis and dark respiration, and distribution of ¹⁴C were determined for new (current season's) and old (previous season's) needles at 10 times during the seasonal development of young Pinus resinosa Ait. trees. The seasonal changes in these factors associated with the development of the new shoot were related to known seasonal patterns of wood formation.

Net photosynthesis per gram of needle dry weight (photosynthetic efficiency) was maximum in the old needles at the time of first new needle elongation; at the same time translocation of ¹⁴C from old to new needles was greatest. Photosynthetic efficiency of new needles was maximum at the end of the period of rapid new needle elongation, when the new needles also began exporting much greater quantities of ¹⁴C to other plant parts. In particular, the amount translocated from the new needles to the stem was greatly increased. At this time thick-walled xylem cells were first observed in the stem.

These results, together with those of previous studies, indicate that the production of thick-walled xylem tracheids normally associated with latewood is physiologically correlated with maturation of the current season's needles. Because there is a lesser demand for photosynthate in the new shoot and a high rate of photosynthesis in the whole plant at the time of new needle maturity, a sharply increased amount of photosynthate becomes available for wall synthesis by cambial derivatives.

     

Physiology of Pinus strobus L. Seedlings Grown under High or Low Soil Moisture Conditions

White pine seedlings (Pinus strobus L.) were grown under highor low soil-moisture levels. The increase in the length andin the fresh weight of seedlings, respiration, photosynthesis,transpiration, translocation of photosynthate from shoots toroots, and bio-electric potentials between the tip and the baseof a stem were measured throughout the growing season from Aprilto October 1964. At both moisture levels the lowest translocation of recent photosynthatefrom shoots to roots occurred during early summer, or at thetime when the rate of root growth was the lowest and that ofthe shoot the highest. The specific activity of ¹⁴CO₂, respiredby the shoots of such plants remained high throughout the 8h of the experiment, indicating a continuous utilization ofrecent photosynthate as a respiratory substrate. On the otherhand, early and late in the growing season, when translocationof recent photosynthate from shoots to roots and the rate ofroot growth were high, the specific activity of ¹⁴CO₂, respiredby the shoots rapidly decreased during the 8 h of the experiment,indicating a drop in the utilization of recent photosynthateas respiratory substrate. The highest positive values for thepotential difference between the top and the base of the mainshoot also occurred in early summer or during the period ofhigh rates of transpiration per needle stomatal surface area.