13C labelling reveals different contributions of photoassimilates from infructescences for fruiting in two temperate forest tree species

The pathways of currently fixed carbon in fruit bearing branchlets were investigated in two temperate forest tree species (CARPINUS BETULUS and FAGUS SYLVATICA), which differ in texture of their vegetative infructescence tissues (leaf-like in CARPINUS vs. woody in FAGUS). During late spring, (13)C pulse-labelling was conducted on girdled, defoliated, girdled plus defoliated and untreated fruiting branchlets of mature trees IN SITU, to assess changes in C relations in response to the introduced C source-sink imbalances. At harvest in early August, 75 - 100 % of the recovered (13)C label was bound to infructescences (either fruits or vegetative infructescence tissue), revealing them as the prime C sinks for current photoassimilates. Leaves on girdled branchlets were not stronger labelled than on ungirdled ones in both species, indicating no upregulation of the leaves' photosynthetic capacity in response to the prevention of phloemic transport, which was also supported by measurements of light saturated photosynthesis. In contrast, (13)C labels tended to be higher after complete defoliation in the vegetative infructescence tissues of CARPINUS, suggesting enhanced net photosynthesis of green infructescence parts as compensation for the loss of regular leaves. The total labelling-derived (13)C content of whole infructescences was very similar between foliated and defoliated CARPINUS branchlets. Cupulae of FAGUS, on the other hand, remained almost unlabelled on defoliated branchlets, indicating the photosynthetic inactivity of this woody infructescence tissue. Consequently, CARPINUS still produced relatively high fruit masses on girdled plus defoliated branchlets, while in FAGUS fruit development ceased almost completely at this most severe treatment. Our results highlight that green vegetative infructescence tissue assimilates substantial amounts of C and can partly substitute regular leaves as C sources for successful fruit development.     

Drought and shade deplete nonstructural carbohydrate reserves in seedlings of five temperate tree species

Plants that store nonstructural carbohydrates (NSC) may rely on carbon reserves to survive carbon‐limiting stress, assuming that reserves can be mobilized. We asked whether carbon reserves decrease in resource stressed seedlings, and if NSC allocation is related to species' relative stress tolerances. We tested the effects of stress (shade, drought, and defoliation) on NSC in seedlings of five temperate tree species (Acer rubrum Marsh., Betula papyrifera Marsh., Fraxinus americana L., Quercus rubra L., and Quercus velutina Lam.). In a greenhouse experiment, seedlings were subjected to combinations of shade, drought, and defoliation. We harvested seedlings over 32–97 days and measured biomass and NSC concentrations in stems and roots to estimate depletion rates. For all species and treatments, except for defoliation, seedling growth and NSC accumulation ceased. Shade and drought combined caused total NSC decreases in all species. For shade or drought alone, only some species experienced decreases. Starch followed similar patterns as total NSC, but soluble sugars increased under drought for drought‐tolerant species. These results provide evidence that species deplete stored carbon in response to carbon limiting stress and that species differences in NSC response may be important for understanding carbon depletion as a buffer against shade‐ and drought‐induced mortality.     

Effects of defoliation and girdling on fruit production in Ilex aquifolium

1. The effects of defoliation and girdling on fruit production in European holly Ilex aquifolium were studied in a northern Spanish population. Three defoliation treatments (control, 50% and 100% leaf removal) were applied at branch level in 10 trees. Six branches were used per treatment in each tree and half of these branches were girdled (a ring of bark and cambium was removed from the branch base).
2. Leaf removal from ungirdled branches had little effect on fruit set, fruit production and reproductive allocation at branch level. However, these variables decreased as the extent of defoliation increased on girdled branches.
3. Fruit production in undefoliated branches did not differ between girdled and ungirdled branches, showing that girdled branches were autonomous for fruit production when undamaged. Mean fruit mass of girdled-100% defoliated branches averaged 8% of fruit mass produced by control branches.
4. Fruit production of ungirdled-100% defoliated branches did not differ from control branches, showing the importance of resource importation from other branches. Hence the branches may be regarded as only partially autonomous for fruit production.
5. Fruit production of girdled-50% defoliated branches was 42% of fruit production of control branches, while ungirdled-50% defoliated branches amounted to 70% of controls. These results also demonstrated resource importation but did not support the existence of photosynthetic compensatory mechanisms.
6. The ability of resource importation developed by fruiting branches might be a mechanism to reduce the effect of reproductive costs in this species.     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

Translocation of carbohydrates and proline in young grapefruit trees at low temperatures

Girdling and defoliation of fruit-bearing grapefruit (Citrus paradisi Macf.) branches inhibited the accumulation of soluble carbohydrates and proline in fruit tissues during low temperature treatment of trees. These treatments did not inhibit hydrolysis of sucrose to reducing sugars. Flavedo and albedo tissues responded similarly to low temperatures but little or no change occurred in the juice. Therefore, soluble carbohydrates and proline do not appear to interchange between different tissues of the fruit at low temperatures but instead are translocated into the fruit from other parts of the plant. Girdling fruit-bearing branches immediately after low temperature treatments inhibited the accumulation of sucrose in fruit tissues at dehardening temperatures. Also, proline levels decreased rapidly in fruit on girdled branches at dehardening temperatures. This rapid decrease suggests proline may serve as a source for respiratory energy in grapefruit during rapidly changing temperatures that favor active growth and during recovery of citrus from environmental stress.     

Fruit production in three masting tree species does not rely on stored carbon reserves

Fruiting is typically considered to massively burden the seasonal carbon budget of trees. The cost of reproduction has therefore been suggested as a proximate factor explaining observed mast-fruiting patterns. Here, we used a large-scale, continuous ¹³C labeling of mature, deciduous trees in a temperate Swiss forest to investigate to what extent fruit formation in three species with masting reproduction behavior (Carpinus betulus, Fagus sylvatica, Quercus petraea) relies on the import of stored carbon reserves. Using a free-air CO₂ enrichment system, we exposed trees to ¹³C-depleted CO₂ during 8 consecutive years. By the end of this experiment, carbon reserve pools had significantly lower δ¹³C values compared to control trees. δ¹³C analysis of new biomass during the first season after termination of the CO₂ enrichment allowed us to distinguish the sources of built-in carbon (old carbon reserves vs. current assimilates). Flowers and expanding leaves carried a significant ¹³C label from old carbon stores. In contrast, fruits and vegetative infructescence tissues were exclusively produced from current, unlabeled photoassimilates in all three species, including F. sylvatica, which had a strong masting season. Analyses of δ¹³C in purified starch from xylem of fruit-bearing shoots revealed a complete turn-over of starch during the season, likely due to its usage for bud break. This study is the first to directly demonstrate that fruiting is independent from old carbon reserves in masting trees, with significant implications for mechanistic models that explain mast seeding.     

Defoliation and below-ground herbivory in the grass Muhlenbergiaquadridentata : Effects on plant performance and on the root-feeder Phyllophaga sp. (Coleoptera, Melolonthidae)

In this study we evaluated (1) the combined effects of simulated defoliation and below-ground herbivory (BGH) on the biomass and nitrogen content of tillers and roots of the bunchgrass Muhlenbergia quadridentata and (2) the effect of defoliation on the survival of third-instar root-feeder larvae of Phyllophaga sp. The experiment was performed in a pine forest area at an altitude of 3200 m above sea level. The grass and the root-feeder species were native and dominant in the understory and in the macroarthropod root-feeder communities, respectively. Plants were established in pots in the field and were subjected to the following treatments in a factorial design: simulated defoliation (three levels) and BGH (with or without root-feeder larvae) with ten replicates per treatment. Plants were defoliated three times at 2-month intervals. The interaction between defoliation and root herbivory was significant for all components of plant biomass. In every case, light defoliation with BGH decreased live above-ground, root and total plant biomass, and the number of live tillers by more than 50% with respect to the same defoliation level without root-feeders. Plants apparently did not compensate for the carbon drain by root-feeders when a high proportion of older leaves were not removed by defoliation. Plants under heavy defoliation were not affected by the presence of root-feeders and showed a greater live/dead above-ground biomass ratio than lightly defoliated and control plants. Defoliation and BGH did not change tiller and root N concentrations but root herbivores did decrease live-tiller N content in lightly defoliated plants. Root-feeders but not defoliation decreased the root/shoot ratio by 40% and the live/dead above-ground biomass ratio by 45% through increased tiller mortality. Survivorship and final biomass of Phyllophaga sp. larvae were not affected by defoliation treatments during the 6-month study period. Received: 17 May 1996 / Accepted: 1 November 1996     

环剥和去叶对红松雌球果和枝生长及不同组织和器官中养分含量的影响

为探索红松雌球果发育和新枝生长的养分来源和供需关系,对生殖母枝进行环剥、去叶及两者相结合处理,研究不同处理对雌球果发育和枝生长及不同组织和器官中碳水化合物(NSC)、氮(N)和磷(P)含量的影响.结果表明:环剥处理对雌球果发育和新枝生长及不同组织和器官中NSC、N和P含量影响显著,去叶处理的影响较小.环剥后母枝木质部和韧皮部NSC含量显著低于对照(CK,不环剥+0%去叶),并随去叶程度的增加显著减少,尤其是环剥+100%去叶的木质部和韧皮部中NSC含量分别比CK减少59.0%和64.8%,NSC的不足导致母枝、新枝死亡和雌球果败育.环剥处理下,0%、50%和100%去叶处理的母枝木质部和韧皮部中N和P含量显著高于CK,其中木质部中N含量比CK分别高17.3%、18.2%和24.3%,P含量比CK分别高17.9%、7.1%和3.6%,韧皮部中N含量比CK分别高39.3%、35.2%和48.9%,P含量比CK分别高31.0%、28.2%和14.8%.红松雌球果发育和新枝生长消耗大量的碳水化合物、N和P,母枝本身制造或贮存的碳水化合物和矿质养分不能满足雌球果发育和新枝生长的需求,碳水化合物和矿质养分需要从母枝以外的其他组织输入.     

干旱-高钙对麻栎幼苗非结构性碳水化合物含量和分配的影响

土壤高钙胁迫是干旱-半干旱区影响树木生长的重要环境因子,为阐明干旱-高钙对树木非结构性碳水化合物(Non-structural carbohydrate,NSC)的含量和分配的影响,以麻栎幼苗为研究对象,阐明干旱和干旱-高钙条件对其生长、光合特征及非结构碳水化合物含量与分配的影响。结果表明:干旱显著降低麻栎幼苗生物量,而干旱-高钙处理较干旱进一步降低了麻栎生物量;干旱-高钙在处理初期就能显著抑制麻栎幼苗净光合速率,处理3个月后干旱和干旱-高钙处理的麻栎幼苗光合速率均显著低于对照;干旱处理麻栎幼苗平均非结构性碳水化合物含量增加19.90%,干旱-高钙处理麻栎幼苗整株的平均NSC含量则显著降低25.62%;干旱和干旱-高钙对麻栎幼苗NSC在不同器官间分配也产生不同影响,干旱条件下麻栎幼苗茎中NSC含量增加最多,较对照增加了52.34%,且淀粉的增高幅度(61.94%)高于可溶性糖(25.53%),干旱、高钙共同作用下麻栎幼苗全株平均NSC含量显著减少的同时,NSC积累在叶中,叶NSC含量显著提高32.31%,根、茎中NSC含量则分别显著降低了49.38%和35.31%。干旱-高钙胁迫降低麻栎幼苗NSC含量,且会减少NSC向枝干和根系分配。     

Vitis vinifera root and leaf metabolic composition during fruit maturation: implications of defoliation

Grapevine (Vitis vinifera) roots and leaves represent major carbohydrate and nitrogen (N) sources, either as recent assimilates, or mobilized from labile or storage pools. This study examined the response of root and leaf primary metabolism following defoliation treatments applied to fruiting vines during ripening. The objective was to link alterations in root and leaf metabolism to carbohydrate and N source functioning under conditions of increased fruit sink demand. Potted grapevine leaf area was adjusted near the start of véraison to 25 primary leaves per vine compared to 100 leaves for the control. An additional group of vines were completely defoliated. Fruit sugar and N content development was assessed, and root and leaf starch and N concentrations determined. An untargeted GC/MS approach was undertaken to evaluate root and leaf primary metabolite concentrations. Partial and full defoliation increased root carbohydrate source contribution towards berry sugar accumulation, evident through starch remobilization. Furthermore, root myo‐inositol metabolism played a distinct role during carbohydrate remobilization. Full defoliation induced shikimate pathway derived aromatic amino acid accumulation in roots, while arginine accumulated after full and partial defoliation. Likewise, various leaf amino acids accumulated after partial defoliation. These results suggest elevated root and leaf amino N source activity when leaf N availability is restricted during fruit ripening. Overall, this study provides novel information regarding the impact of leaf source restriction, on metabolic compositions of major carbohydrate and N sources during berry maturation. These results enhance the understanding of source organ carbon and N metabolism during fruit maturation.     

Effect of different leaf-to-fruit ratios on photosynthesis and fruit growth in olive (Olea europaea L.)

The influence of different leaf-to-fruit (l-t-f) ratios on leaf net photosynthetic rate (P _N) and fruit characteristics in Olea europaea L. cv. Frantoio was evaluated in 2001 and 2002. In both years, at the end of June, at the end of July, and in mid-September (first, second, and third time of treatment, respectively), defoliation or fruit thinning were performed to give l-t-f ratios of 1/1, 3/1, 5/1, and 7/1 (about 5.1, 15.3, 25.6, and 35.8 cm² of leaf area per fruit, respectively) on girdled and ungirdled peripheral shoots. P _N showed substantial seasonal and diurnal variations. In ungirdled shoots, no differences due to the different l-t-f ratios were observed, whereas in girdled shoots P _N tended to be lower in shoots with a high l-t-f ratio. In general, the values of leaf transpiration rate (E), stomatal conductance (g _s), sub-stomatal CO₂ concentration (C _i), and dark respiration rate (R _D) were associated with those of P _N. The starch and reducing sugar contents and area leaf dry mass (ADM) tended to be higher in leaves on girdled shoots with high l-t-f ratio, whereas in ungirdled shoots no differences related to the different l-t-f ratios were observed. The higher saccharide content in the leaves and the lower P _N, in the presence of a high C _i, observed in girdled shoots with a high l-t-f ratio suggests that the depression in P _N in these shoots may be the result of a feedback inhibition of the photosynthetic mechanism that regulates such a process. The l-t-f ratio did not have a substantial effect on fruit drop. In ungirdled shoots, the different l-t-f ratios did not produce significant differences in terms of fruit growth and leaf dry matter and saccharide contents, whereas in girdled shoots fruit growth increased as the l-t-f ratio increased, particularly when treatments were applied at the initial stage of fruit development. The percentage of oil in the pulp, on a dry matter basis, was not substantially influenced by girdling and l-t-f ratio. The abundant availability of assimilates seemed to cause earlier fruit ripening and, at the same time, retard fruit senescence (fruit detachment force). Shoot growth was slightly reduced by girdling. The abundant availability of assimilates, induced by girdling associated with high l-t-f ratio, stimulated flower induction.     

Extreme defoliation reduces tree growth but not C and N storage in a winter-deciduous species

Background and Aims There is a growing concern about how forests will respond to increased herbivory associated with climate change. Carbon (C) and nitrogen (N) limitation are hypothesized to cause decreasing growth after defoliation, and eventually mortality. This study examines the effects of a natural and massive defoliation by an insect on mature trees’ C and N storage, which have rarely been studied together, particularly in winter-deciduous species.Methods Survival, growth rate, carbon [C, as non-structural carbohydrate (NSC) concentration] and nitrogen (N) storage, defences (tannins and total polyphenols), and re-foliation traits were examined in naturally defoliated and non-defoliated adult trees of the winter-deciduous temperate species Nothofagus pumilio 1 and 2 years after a massive and complete defoliation caused by the caterpillar of Ormiscodes amphimone (Saturniidae) during summer 2009 in Patagonia.Key Results Defoliated trees did not die but grew significantly less than non-defoliated trees for at least 2 years after defoliation. One year after defoliation, defoliated trees had similar NSC and N concentrations in woody tissues, higher polyphenol concentrations and lower re-foliation than non-defoliated trees. In the second year, however, NSC concentrations in branches were significantly higher in defoliated trees while differences in polyphenols and re-foliation disappeared and decreased, respectively.Conclusions The significant reduction in growth following defoliation was not caused by insufficient C or N availability, as frequently assumed; instead, it was probably due to growth limitations due to factors other than C or N, or to preventative C allocation to storage. This study shows an integrative approach to evaluating plant growth limitations in response to disturbance, by examining major resources other than C (e.g. N), and other C sinks besides storage and growth (e.g. defences and re-foliation).     

去叶对青杨雌雄植株生长和非结构性碳水化合物的影响

近年来,森林食叶害虫在全世界呈爆发趋势.树木的非结构性碳水化合物(NSC)如何响应叶片损失对其生长和生存至关重要.雌雄异株植物在维持森林生态系统稳定性方面扮演着重要角色.然而,目前对该类植物性别之间如何响应去叶的研究还比较少.本文以我国重要的经济和生态恢复树种青杨(Populus cathayana)为研究材料,比较了...     

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves

We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species –Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina– with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots. Received: 14 December 1999 / Accepted: 2 June 1999     

Grain Amaranths Are Defoliation Tolerant Crop Species Capable of Utilizing Stem and Root Carbohydrate Reserves to Sustain Vegetative and Reproductive Growth after Leaf Loss

Tolerance to defoliation can be defined as the degree to which productivity is affected by photosynthetic area reduction. This trait was studied in grain amaranth (Amaranthus cruentus and A. hypochondriacus), which are considered to be a highly defoliation-tolerant species. The physiological and biochemical responses to increasing levels of mechanical leaf removal up to total defoliation were quantified. Tolerance appeared to be dependent on various factors: ( i) amount of lost tissue; (ii) mechanics of leaf tissue removal; (iii) environment, and (iv) species tested. Thus, grain amaranth was found to be a highly tolerant species under green-house conditions when leaf tissue loss was performed by gradual perforation. However, tolerance was compromised under similar conditions when defoliation was done by gradual cutting of the leaf. Also tolerance in completely defoliated plants tended to decrease under field conditions, where differences between A. cruentus and A. hypochondriacus were observed. All non-structural carbohydrate (NSC) levels were reduced in stems and roots of totally defoliated amaranths one day after treatment. Such depletion probably provided the carbon (C) resources needed to sustain the early recovery process in the absence of photosynthetic capacity. This was corroborated by shading of intact plants, which produced the same rapid and drastic reduction of NSC levels in these tissues. These results emphasize the role of stored NSCs, particularly starch, in buffering the impact of severe defoliation in amaranth. The fall in sucrose synthase and cell wall invertase activity observed in stems and roots soon after defoliation was consistent with their predicted shift from sink to source tissues. It is concluded that mobilization of C stores in stems and roots, is a physiologically important trait underlying tolerance to defoliation in grain amaranth.     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Non-structural carbohydrate pools in a tropical forest

The pool size of mobile, i.e. non-structural carbohydrates (NSC) in trees reflects the balance between net photosynthetic carbon uptake (source) and irreversible investments in structures or loss of carbon (sink). The seasonal variation of NSC concentration should reflect the sink/source relationship, provided all tissues from root to crown tops are considered. Using the Smithsonian canopy crane in Panama we studied NSC concentrations in a semi-deciduous tropical forest over 22 months. In the 9 most intensively studied species (out of the 17 investigated), we found higher NSC concentrations (starch, glucose, fructose, sucrose) across all species and organs in the dry season than in the wet season (NSC 7.2% vs 5.8% of dry matter in leaves, 8.8/6.0 in branches, 9.7/8.5 in stems, 8.3/6.4 in coarse and 3.9/2.2 in fine roots). Since this increase was due to starch only, we attribute this to drought-constrained growth (photosynthesis less affected by drought than sink activity). Species-specific phenological rhythms (leafing or fruiting) did not overturn these seasonal trends. Most of the stem volume (diameter at breast height around 40 cm) stores NSC. We present the first whole forest estimate of NSC pool size, assuming a 200 t ha^–1 forest biomass: 8% of this i.e. ca. 16 t ha^–1 is NSC, with ca. 13 t ha^–1 in stems and branches, ca. 0.5 and 2.8 t ha^–1 in leaves and roots. Starch alone (ca. 10.5 t ha^–1) accounts for far more C than would be needed to replace the total leaf canopy without additional photosynthesis. NSC never passed through a period of significant depletion. Leaf flushing did not draw heavily upon NSC pools. Overall, the data imply a high carbon supply status of this forest and that growth during the dry season is not carbon limited. Rather, water shortage seems to limit carbon investment (new tissue formation) directly, leaving little leeway for a direct CO₂ fertilization effects.     

THE POLLINATION BIOLOGY OF TILIA

An anthecological study of Tilia in the Great Plains and New England was undertaken for both native and introduced species. The floral bracts are postulated as being at least as important in pollinator attraction as they are in fruit dispersal. A characteristic sweet odor is always present, but becomes stronger at dusk. Flowers are protandrous with anthers opening for the first time late in the afternoon and releasing pollen abundantly for about 24 hr. Initial nectar production coincides with stigma receptivity which begins late in the afternoon, but on the second day a flower is open. Sixty-six species of insects in 29 families were collected. Bees and flies are the most common diurnal visitors and moths are the primary nocturnal visitors. Samples of pollen taken from the insects indicate a relatively high constancy. Experimental tests show that Tilia is not apomictic or self compatible. Anemophily plays a secondary role to entomophily in pollination. Nocturnal pollinators are slighly less effective than diurnal pollinators in effecting fruit set. Although nocturnal pollinators are favored by the syndrome of floral characteristics, it is concluded that Tilia is a generalist in regard to pollination. It is proposed that the lack of clear morphological differences between Tilia species leads to pollinator inconstancy and thus hybridization.     

Fruit Production in Mature and Recently Regenerated Forests of the Appalachians

ABSTRACT Fleshy fruit is a key food resource for both game and nongame wildlife, and it may be especially important for migratory birds during fall and for resident birds and mammals during winter. Land managers need to know how land uses affect the quantities and species of fruit produced in different forest types and how fruit production varies seasonally and as young stands mature. During June 1999-April 2004, we quantified fleshy fruit abundance monthly in 31 0.1-ha plots in 2 silvicultural treatments: 1) young 2-age stands with low basal area retention, created by shelterwood-with-reserves regeneration cuts (R; harvested 1998–1999); and 2) uncut mature closed-canopy stands (M) in 2 common southern Appalachian, USA, forest types (upland hardwood and cove hardwood [CH] forests). Over the 5-year study period, total dry pulp biomass production was low and relatively constant in both M forest types (x̄ = 0.5-2.0 kg/ha). In contrast, fruit production increased each year in R, and it was 5.0 to 19.6 times greater in R than in M stands beginning 3–5 years postharvest. Two disturbance-associated species, pokeweed (Phytolacca americana) and blackberry (Rubus allegheniensis), produced a large proportion of fruit in R but showed different patterns of establishment and decline. Huckleberry (Gaylussacia ursina) recovered rapidly after harvest and was a major producer in both silvicultural treatments and forest types each year. Several herbaceous species that are not associated with disturbance produced more fruit in CHR. Few species produced more fruit in M than in R. Fruit production by most tree species was similar between R and M, due to fruiting by stump sprouts in R within 1–3 years postharvest. Fruit availability was highest during summer and early fall. American holly (Ilex opaca), sumac (Rhus spp.), and greenbriar (Smilax spp.) retained fruit during winter months but were patchy in distribution. In the southern Appalachians, young recently regenerated stands provide abundant fruit compared to mature forest stands and represent an important source of food for wildlife for several years after harvest. Fruit availability differs temporally and spatially because of differences in species composition, fruiting phenology, and the dynamic process of colonization and recovery in recently harvested stands. Land managers could enhance fruit availability for many game and nongame species by creating or maintaining young stands within forests.     

Mycorrhiza and seedling establishment in a subarctic meadow: Effects of fertilization and defoliation

Abstract. Question: How does changing resource availability induced by fertilization and defoliation affect seedling establishment and mycorrhizal symbiosis in a subarctic meadow? Location: 610 m a.s.l., Kilpisjärvi (69°03’N, 20°50’E), Finland. Methods: A short‐term full‐factorial experiment was established, with fertilization and defoliation of natural established vegetation as treatments. Seeds of two perennial herbs Solidago virgaurea and Gnaphalium norvegicum were sown in natural vegetation and their germination and growth followed. At the final harvest we measured the response in terms of arbuscular mycorrhizal (AM) colonization, biomass and nitrogen concentration of the seedlings and the established vegetation. Results: Germination rate was negatively affected by defoliation in the unfertilized plots. The shoot biomass of S. virgaurea seedlings was reduced by the defoliation and fertilization treatments, but not affected by their interaction. In G. norvegicum, the germination rate and the seedling shoot biomass were negatively correlated with moss biomass in the plots. In the established plants the arbuscular colonization rate was low and defoliation and fertilization treatments either increased or did not affect the colonization by AM fungi. In the seedlings, the colonization rate by AM fungi was high, but it was not affected by treatments. Both seedlings and established plants were colonized by dark septate fungi. Conclusions: Reduction of plant biomass by herbivores can have different effects on seedling growth in areas of high and low soil nutrient availability. The weak response of AM colonization to defoliation and fertilization suggests that AM symbiosis is not affected by altering plant resource availability under the conditions employed in this study.