Corticular Photosynthesis Research Articles

Possible Contribution of Corticular Photosynthesis to Grapevine Winter Hardiness

Numerous studies show that photosynthesis in non-foliar tissues contributes to plant productivity. Here, we demonstrate that in chlorenchyma tissues of lignified branches of grape vines, photosynthetic activity is maintained during winter and provide evidence that corticular photosynthesis could contribute to the plant’s freeze tolerance. In a collection of grape varieties that varied noticeably in freeze tolerance, a positive correlation between the maximum quantum yield of photosystem II in the wintering vines and the ability to survive harsh winter temperatures was observed. A more detailed comparison of two grapevine varieties differing in freeze tolerance showed that the vines of the more tolerant variety have more abundant corticular chlorenchyma with chloroplasts containing a better developed network of photosynthetic membranes, characterized by a higher photosynthetic pigments content, higher efficiency of both photosystems, and higher mobility of antennae complexes under the changing light intensity. In addition, we found that freezing temperatures induced more damage in vine samples when they were preliminarily treated with a specific inhibitor of photosynthetic electron transfer. The data obtained could be useful in the generation of freeze-tolerant grape varieties.

Can stress turn trees hair white? Hair covering of stems improves resilience of corticular photosynthesis against heat-stress

The hypothesis was tested that hair covering of stems improves resilience of corticular photosynthesis against heat stress. Hairy and non-hairy outer bark of Quercus ilex L. and Quercus robur L. trees was removed and optical properties measured. Additionally, structural bark traits and chlorophyll fluorescence parameters during heat stress treatment were studied. Optical analysis revealed a protective role of hairy outer bark (OB) against overheating of the underlying cortex of the stems. Hairiness decreased OB transmittance and increased thermal insulation of stems by an increased absorptance and reflectance of OB in the visible (380–720 nm) and an increased reflectance in the infrared part of the spectrum (720–900 nm). Simple linear regression analysis revealed no significant effect of stem structural traits (OB thickness (OBT), cortex density (Dcortex), cortex water content (Wcortex)) on corticular photochemistry (PScort), while optical traits of outer bark were significantly (P < 0.01) correlated with PScort. OB reflectance explained up to 91% of the variation in PSII quantum yield under heat stress. At high temperatures (> 45 °C) PScort of the hairy species showed a higher resilience and a better post-stress-recovery as compared to the non-hairy one. It is concluded that stem hairs play a physiologically significant role in modulating the stem energy balance due to a close interaction between optical characteristics of hairy OB and stem photochemical processes.

Involvement of stem corticular photosynthesis in hydraulic maintenance of Eucalyptus trees and its effect on leaf gas exchange

Different from leaf photosynthesis that depends on stomatal uptake of CO2, stem green tissue mainly uses internal CO2 for re-assimilation. Research attention has moved to its specific effect on xylem hydraulic function, however, the effect of stem corticular photosynthesis on whole-plant water transport and leaf photosynthesis during drought stress of varying intensities remains unexplored. To fill this knowledge gap, we designed stem shading treatments on Eucalyptus urophylla trees in the field and its seedlings in the greenhouse with soil water-controlled experiment. Sap flow density (Js), whole-plant hydraulic conductance (Kplant), stem tissue fluorescence parameters and stem non-structural carbohydrates (NSC) were measured. Stem shading treatments caused the decrease of Kplant and Js of E. urophylla trees, more importantly, the increase of Kplant and Js with the recovery of corticular photosynthesis after shading removal was observed in the field experiment. The main effect of shading treatment on stem NSC concentration of E. urophylla seedlings is a decrease, which was not influenced by soil water conditions. Moreover, the shading effects on leaf gas exchange vary with drought intensity, under severe drought stress, stem shading caused a significant decline of stomatal conductance (gs,) and leaf maximum net photosynthesis (Amax) for E. urophylla seedlings. These findings suggested that stem corticular photosynthesis plays a compensatory role in maintaining whole-plant hydraulic conductance integrity and water transport of E. urophylla. The remarkable negative impact of stem shading on leaf gas exchange implying a possible interconnection between leaf and stem photosynthesis, which highlights the potential significance of stem photosynthesis in drought stress resistance under changing climate regimes.

Corticular photosynthesis drives bark water uptake to refill embolized vessels in dehydrated branches of Salix matsudana.

It is well known that xylem embolism can be repaired by bark water uptake and that the sugar required for embolism refilling can be provided by corticular photosynthesis. However, the relationship between corticular photosynthesis and embolism repair by bark water uptake is still poorly understood. In this study, the role of corticular photosynthesis in embolism repair was assessed using Salix matsudana branch segments dehydrated to -1.9MPa (P50 , water potential at 50% loss of conductivity). The results indicated that corticular photosynthesis significantly promoted water uptake and nonstructural carbohydrate (NSC) accumulation in the bark and xylem during soaking, thereby effectively enhancing the refilling of the embolized vessels and the recovery of hydraulic conductivity. Furthermore, the influence of the extent of dehydration on the embolism refilling enhanced by corticular photosynthesis was investigated. The enhanced refilling effects were much higher in the mildly dehydrated (-1.5MPa) and moderately dehydrated (-1.9MPa) branch segments than in the severely dehydrated (-2.2MPa) branch segments. This study provides evidence that corticular photosynthesis plays a crucial role in xylem embolism repair by bark water uptake for mildly and moderately dehydrated branches.

More than just CO2 -recycling: corticular photosynthesis as a mechanism to reduce the risk of an energy crisis induced by low oxygen.

Reassimilation of internal CO2 via corticular photosynthesis (PScort ) has an important effect on the carbon economy of trees. However, little is known about its role as a source of O2 supply to the stem parenchyma and its implications in consumption and movement of O2 within trees. PScort of young Populus nigra (black poplar) trees was investigated by combining optical micro-optode measurements with monitoring of stem chlorophyll fluorescence. During times of zero sap flow in spring, stem oxygen concentrations (cO2 ) exhibited large temporal changes. In the sapwood, over 80% of diurnal changes in cO2 could be explained by respiration rates (Rd(mod) ). In the cortex, photosynthetic oxygen release during the day altered this relationship. With daytime illumination, oxygen levels in the cortex steadily increased from subambient and even exhibited a diel period of superoxia of up to 110% (% air sat.). By contrast, in the sapwood, cO2 never reached ambient levels; the diurnal oxygen deficit was up to 25% of air saturation. Our results confirm that PScort is not only a CO2 -recycling mechanism, it is also a mechanism to actively raise the cortical O2 concentration and counteract temporal/spatial hypoxia inside plant stems.

Tree Species with Photosynthetic Stems Have Greater Nighttime Sap Flux.

An increasing body of evidence has shown that nighttime sap flux occurs in most plants, but the physiological implications and regulatory mechanism are poorly known. The significance of corticular photosynthesis has received much attention during the last decade, however, the knowledge of the relationship between corticular photosynthesis and nocturnal stem sap flow is limited at present. In this study, we divided seven tree species into two groups according to different photosynthetic capabilities: trees of species with (Castanopsis hystrix, Michelia macclurei, Eucalyptus citriodora, and Eucalyptus grandis × urophylla) and without (Castanopsis fissa, Schima superba, and Acacia auriculiformis) photosynthetic stems, and the sap flux (Js) and chlorophyll fluorescence parameters for these species were measured. One-way ANOVA analysis showed that the Fv/Fm (Maximum photochemical quantum yield of PSII) and ΦPSII (effective photochemical quantum yield of PSII) values were lower in non-photosynthetic stem species compared to photosynthetic stem species. The linear regression analysis showed that Js,d (daytime sap flux) and Js,n (nighttime sap flux) of non-photosynthetic stem species was 87.7 and 60.9% of the stem photosynthetic species. Furthermore, for a given daytime transpiration water loss, total nighttime sap flux was higher in species with photosynthetic stems (SlopeSMA = 2.680) than in non-photosynthetic stems species (SlopeSMA = 1.943). These results mean that stem corticular photosynthesis has a possible effect on the nighttime water flow, highlighting the important eco-physiological relationship between nighttime sap flux and corticular photosynthesis.

Photosynthetic refixation varies along the stem and reduces CO2 efflux in mature boreal Pinus sylvestris trees.

Trees are able to reduce their carbon (C) losses by refixing some of the CO2 diffusing out of their stems through corticular photosynthesis. Previous studies have shown that under ideal conditions the outflowing CO2 can be completely assimilated in metabolically active, young stem and branch tissues. Fewer studies have, however, been carried out on the older stem sections of large trees and, accordingly, the importance of refixation is still unclear under natural environmental conditions. We investigated the spatial and temporal variation in refixation in ~90-year-old boreal Scots pine (Pinus sylvestris L.) trees by utilizing month-long continuous measurements of stem CO2 efflux (Ec) made at four heights along the bole. Refixation rates were found to vary considerably along the bole, leading to a 28% reduction in long-term Ec in the upper stem compared with a negligible reduction at breast height. This vertical pattern correlated with variation in light availability, bark chlorophyll content and bark type. Analysis of the vertical and diurnal patterns in Ec further suggested that the influence of sap flow on the observed daytime reduction in Ec was small. The areal rates of corticular photosynthesis were much lower than previous estimates of photosynthetic rates per unit leaf area from the same trees, implying that the impact of refixation on tree-scale C uptake was small. However, upscaling of refixation indicated that 23-27% of the potential Ec was refixed by the bole and the branches, thereby significantly reducing the woody tissue C losses. Thus, our results suggest that refixation needs to be considered when evaluating the aboveground C cycling of mature P. sylvestris stands and that breast-height estimates should not be extrapolated to the whole tree.

树干皮层光合作用：生理生态功能和测定方法

树干皮层光合作用：生理生态功能和测定方法

Chlorophyll fluorescence for visualizing the spatial and temporal spread of Phytophthora alni subsp. alni in alder bark tissue

The impact of alder Phytophthora (Phytophthora alni subsp. alni) on corticular photosynthetic metabolism was explored via measurements of chlorophyll fluorescence. Stem inoculation induced a sharp reduction of maximum (Fv/Fm) and effective quantum yield of PSII (). Observations of the axial and radial spread of the pathogen revealed that near to the point of inoculation and in the whole centre of the stem lesion Fv/Fm and of the cortex chlorenchyma decreased to almost zero, indicating tissue necrosis. Low values of Fv/Fm and were also found in some presymptomatic regions beyond the visible stem lesion. In contrast, substantial photosynthetic activity was found in uninvaded parts of the inoculated trees and in the control. These stem parts showed a marked light‐adapted quantum efficiency of PSII as well as marked electron transport rates in their bark tissues. Thus, corticular photosynthesis stayed unaffected in these stem parts supporting stem carbon balance. Chlorophyll fluorescence measurements in the field illustrated that stem infection with P. alni subsp. alni and the effect on the bark tissues is not only highly heterogeneous but also underlies very quick temporal changes, due to a rapid destruction of the photosynthetic apparatus. The results show that chlorophyll fluorescence measurements and fluorescence imaging are useful indicators of tissue infection caused by Phytophthora infection of bark tissues. Chlorophyll fluorescence measurements can be used to map and visualize the spatial as well as temporal spread of bark pathogens and can give a first indication of invasion of the host tissue beyond the visible lesion.

Contribution of stem CO&lt;sub&gt;2&lt;/sub&gt; fixation to whole-plant carbon balance in nonsucculent species

In many plant species that remain leafless part of the year, CO2 fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO2 fixation by stems with stomata in the epidermis and net corticular CO2 fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO2 ling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO2 uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO2 occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO2, the latter figure implying net CO2 uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.

Contribution of corticular photosynthesis to bud development in African baobab (Adansonia digitata L.) and Castor bean (Ricinus communis L.) seedlings

The African baobab (Adansonia digitata L.) and castor bean (Ricinus communis L.) are drought resistant green-stemmed succulent plants which grow in the arid and semi-arid regions of Africa. Photosynthesis in the stems of green-stemmed plants is known to contribute to plant carbon gain especially during leafless periods. To study the contribution of stem photosynthesis in stem succulent plants, the height and stem diameter of baobab and castor bean plants grown in the greenhouse were measured. The plants were completely defoliated and subjected to different treatments: Watered with open stems (WO), watered and stems covered with aluminium foil (WC) to achieve 100% light exclusion, drought and open (DO) and drought and covered (DC). Stem coverage with aluminium foil resulted in a higher stem height and diameter during drought for baobab with similar trends seen in castor bean. Light exclusion resulted in a significantly lower bud DW production and enrichment in 13C in bud dry matter of castor bean and in stem dry matter of baobab. These show that corticular photosynthesis contributes in carbon gain in these species.

Stable isotopes reveal the contribution of corticular photosynthesis to growth in branches of Eucalyptus miniata.

The deciduous bark habit is widespread in the woody plant genus Eucalyptus. Species with deciduous bark seasonally shed a layer of dead bark, thereby maintaining smooth-bark surfaces on branches and stems as they age and increase in diameter. This has a significant cost in terms of fire protection, because smooth-barked species have thinner bark than rough-barked species that accumulate successive layers of dead bark. Eucalypts are closely associated with fire, suggesting that the smooth-bark habit must also provide a significant benefit. We suggest that this benefit is corticular photosynthesis. To test this, we quantified the contribution of corticular photosynthesis to wood production in smooth-barked branches of Eucalyptus miniata growing in tropical savanna in northern Australia. We covered branch sections with aluminum foil for 4 years to block corticular photosynthesis and then compared the oxygen and carbon stable isotope composition of foil-covered and uncovered branch sections. We developed theory to calculate the proportion of wood constructed from corticular photosynthate and the mean proportional refixation rate during corticular photosynthesis from the observed isotopic differences. Coverage with aluminum foil for 4 years increased wood δ(13)C by 0.5‰ (P = 0.002, n = 6) and wood δ(18)O by 0.5‰ (P = 0.02, n = 6). Based on these data, we estimated that 11% ± 3% of wood in the uncovered branch sections was constructed from corticular photosynthate, with a mean δ(13)C of -34.8‰, and that the mean proportional refixation rate during corticular photosynthesis was 0.71 ± 0.15. This demonstrates that corticular photosynthesis makes a significant contribution to the carbon economy of smooth-barked eucalypts.

Role of corticular photosynthesis following defoliation in Eucalyptus globulus

Defoliation can reduce net fixation of atmospheric CO(2) by the canopy, but increase the intensity and duration of photosynthetically active radiation on stems. Stem CO(2) flux and leaf gas exchange in young Eucalyptus globulus seedlings were measured to assess the impact of defoliation on these processes and to determine the potential contribution of re-fixation by photosynthetic inner bark in offsetting the effects of defoliation in a woody species. Pot and field trials examined how artificial defoliation of the canopy affected the photosynthetic characteristics of main stems of young Eucalyptus globulus seedlings. Defoliated potted seedlings were characterized by transient increases in foliar photosynthetic rates and concomitant decreases in stem CO(2) fluxes (both in the dark and light). Defoliated field-grown seedlings showed similar stem CO(2) flux responses, but of reduced magnitude. Despite demonstrating increased re-fixation capability, defoliated potted-seedlings had slowed stem growth. The green stem of seedlings exhibited largely shade-adapted characteristics. Defoliation reduced stem chlorophyll a/b ratio and increased carotenoid concentration. An increased capacity to re-fix internally respired CO(2) (up to 96%) suggested that stem re-fixation represents a previously unexplored mechanism to minimize the impact of foliar loss by maximizing the contribution of all photosynthetic tissues, particularly for young seedlings.

Corticular photosynthesis: A mechanism to enhance shrub expansion in coastal environments

Myrica cerifera L. (Myricaceae), the dominant woody species on many barrier islands along the southeastern coast of the United States, is expanding into grass-dominated, mesic, interdunal depressions where it forms dense thickets. Expansion may be attributed to a symbiotic nitrogen fixation with the bacterium Frankia, an evergreen leaf habit and, possibly, corticular photosynthesis (CP, i.e. refixation of respired CO2, %ref). We quantified seasonal variations in CP characteristics in first through fifth order branches of M. cerifera to determine the extent and relevance of CP to shrub expansion in coastal environments. Maximum mean %ref was 110&plusmn;39 % of CO2 efflux in the dark (R D) in first order branches during winter. Minimum %ref was 18&plusmn;3 % in fifth order branches during summer. Variations in %ref paralleled changes in incident photosynthetic photon flux density (PPFD). As incident PPFD attenuated with increasing branch order, %ref decreased. A less dense canopy in winter led to increased PPFD and increases in %ref. Total chlorophyll (Chl) content and Chl a/b ratios were consistent with shade acclimation as branch order increased. CP may be a mechanism to enhance M. cerifera shrub expansion because of the potential increase in whole plant carbon use efficiency and water use efficiency attributed to refixation of respired CO2.

CO2 efflux, CO2 concentration and photosynthetic refixation in stems of Eucalyptus globulus (Labill.)

In spite of the importance of respiration in forest carbon budgets, the mechanisms by which physiological factors control stem respiration are unclear. An experiment was set up in a Eucalyptus globulus plantation in central Portugal with monoculture stands of 5-year-old and 10-year-old trees. CO(2) efflux from stems under shaded and unshaded conditions, as well as the concentration of CO(2) dissolved in sap [CO(2)(*)], stem temperature, and sap flow were measured with the objective of improving our understanding of the factors controlling CO(2) release from stems of E. globulus. CO(2) efflux was consistently higher in 5-year-old, compared with 10-year-old, stems, averaging 3.4 versus 1.3 mumol m(-2) s(-1), respectively. Temperature and [CO(2)(*)] both had important, and similar, influences on the rate of CO(2) efflux from the stems, but neither explained the difference in the magnitude of CO(2) efflux between trees of different age and size. No relationship was found between efflux and sap flow, and efflux was independent of tree volume, suggesting the presence of substantial barriers to the diffusion of CO(2) from the xylem to the atmosphere in this species. The rate of corticular photosynthesis was the same in trees of both ages and only reduced CO(2) efflux by 7%, probably due to the low irradiance at the stem surface below the canopy. The younger trees were growing at a much faster rate than the older trees. The difference between CO(2) efflux from the younger and older stems appears to have resulted from a difference in growth respiration rather than a difference in the rate of diffusion of xylem-transported CO(2).

General trait relationships in stems: a study on the performance and interrelationships of several functional and structural parameters involved in corticular photosynthesis

We addressed corticular photosynthesis, focusing on parameters of underlying dark and light reactions as well as structural differentiation. To unveil general stem traits and underlying principles that may be valid across several tree species, CO(2) exchange rates and chlorophyll-fluorescence parameters were measured in current-year to 3-year-old stems of five deciduous tree species (including climax and pioneer species). Across species, dark CO(2) efflux rates (R(d)) of stems exhibited a common regression relationship with photosynthetic rates (A) and light-adapted quantum efficiency of photosystem II (PSII) (Delta F/Fm'), a pattern analogous to leaf trait correlations. Furthermore, A and Delta F/Fm' were closely interrelated to each other. Consistent correlations of stem structure and function were also assessed among species. Changes in tissue structure during ageing significantly affected several stem functional parameters. Stem CO(2) efflux during the dark and corticular photosynthetic rates declined with increasing stem age as well as light-adapted quantum efficiency of PSII. Furthermore, a strong relationship between stem R(d) and peridermal PFD-transmittance (T) as well as between R(d) and total bark chlorophyll was evident. Consistent results were found for the relationships between corticular photosynthesis (or primary photosynthetic reactions like Delta F/Fm') and selected structural traits. The found correlation patterns among functional and/or structural traits of stems and their concordance with leaf trait relationships may aid in identifying underlying mechanisms and scaling relationships that link traits to plant and ecosystem function.

Antitranspirant functions of stem periderms and their influence on corticular photosynthesis under drought stress

Transpiration and photosynthesis of current-year stems and adult leaves of different deciduous tree species were investigated to estimate their probable influence on carbon balance. Peridermal transpiration of young stems was found to be rather small as compared to the transpiration of leaves (stem/leaf like 1/5–1/20). A characteristic that was mainly attributable to the lower peridermal conductance to water and CO2, which made up only 8–28% of stomatal conductance. Water vapour conductance was significantly lower in stems, but also non-responsive to PAR, which led to a comparatively higher water use efficiency (WUE, ratio assimilation/transpiration). Thus, although corticular photosynthesis reached only 11–37% of leaf photosynthesis, it may be a means of improving the carbon balance of stems under limited water availability. The influence of drought stress on primary photosynthetic reactions was also studied. Under simulated drought conditions the drying time needed to provoke a 50% reduction (t 50) in dark- and light-adapted PSII efficiency (Fv/Fm, ΔF/Fm′) was up to ten times higher in stems than in leaves. Nevertheless, up to a relative water deficit (RWD) of around 40–50% dark-adapted PSII efficiency of leaves and stems was rather insensitive to dehydration, showing that the efficiency of open PS II reaction centres is not impaired. Thus, it may be concluded that in stems as well as in leaves the primary site of drought damage is at the level of dark enzyme reactions and not within PSII. However, enduring severe drought caused photoinhibitory damage to the photosynthetic apparatus of leaves and stems; thereby RWD50 values (= RWD needed to provoke a 50% reduction in Fv/Fm ad ΔF/Fm′) were comparably lower in stems as compared to leaves, indicating a possibly higher drought sensitivity of the cortex chlorenchyma.

Photosynthetic characteristics of olive tree (Olea europaea) bark

Functional and structural characteristics of corticular photosynthesis of sun-exposed bark of olive tree (Olea europaea L.) were examined. Stomata are only sporadically present during stem primary growth. Light transmission through the phellem was age dependent, decreasing rapidly in stems older than five years of age. Light transmission was also low in pubescent 1-year-old stems. Light transmission was about 50% higher in wet phellem than in dry phellem. Photosynthetic capacity on a unit area basis (measured with an oxygen disc electrode at 27 degrees C and about 5% CO(2) on chlorophyllous tissue discs isolated from the stem) was higher in 1-, 20- and 30-year-old stems compared with 2-10-year-old stems. Low chlorophyll a/b ratio and light compensation points were recorded in olive stems with low phellem light transmission, in accordance with the shade acclimation hypothesis. The intrinsic photochemical efficiency of photosystem II of all stems, especially young stems, was less than that of the leaves. Our results show that olive tree bark possesses an efficient photosynthetic mechanism that may significantly contribute not only to the reduction in concentrations of CO(2) in the inner bark, but also to whole-tree carbon balance.

Effects of ozone impact on the gas exchange and chlorophyll fluorescence of juvenile birch stems ( Betula pendula Roth.)

Effects of ozone impact on gas exchange and chlorophyll fluorescence of juvenile birch ( Betula pendula) stems and leaves were investigated. Significant differences in the response of leaves and stems to ozone were found. In leaves, O 3 exposure led to a significant decline in photosynthetic rates, whereas stems revealed an increased dark respiration and a concomitant increase in corticular photosynthesis. In contrast to birch leaves, corticular photosynthesis appeared to support the carbon balance of stems or even of the whole-tree under O 3 stress. The differences in the ozone-response between leaves and stems were found to be related to ozone uptake rates, and thus to inherent differences in leaf and stem O 3 conductance.

Soil DIC uptake and fixation in Pinus taeda seedlings and its C contribution to plant tissues and ectomycorrhizal fungi

Plants can acquire carbon from sources other than atmospheric carbon dioxide (CO(2)), including soil-dissolved inorganic carbon (DIC). Although the net flux of CO(2) is out of the root, soil DIC can be taken up by the root, transported within the plant, and fixed either photosynthetically or anaplerotically by plant tissues. We tested the ability of Pinus taeda L. seedlings exposed to (13)C-labeled soil DIC and two NH(4)(+) availability regimes to take up and fix soil DIC. We also measured the concentration and distribution of the fixed soil DIC within the plant and mycorrhizal tissues, and quantified the contribution of soil DIC to whole-plant carbon (C) gain. Seedlings exposed to labeled DIC were significantly enriched in (13)C compared with seedlings exposed to unlabeled DIC (6.7 versus -31.7 per thousand). Fixed soil DIC was almost evenly distributed between above- and belowground biomass (55 and 45%, respectively), but was unevenly distributed among tissues. Aboveground, stem tissue contained 65% of the fixed soil DIC but represented only 27% of the aboveground biomass, suggesting either corticular photosynthesis or preferential stem allocation. Belowground, soil DIC had the greatest effect (measured as (13)C enrichment) on the C pool of rapidly growing nonmycorrhizal roots. Soil DIC contributed approximately 0.8% to whole-plant C gain, and approximately 1.6% to belowground C gain. We observed a slight but nonsignificant increase in both relative C gain and the contribution of soil DIC to C gain in NH(4)(+)-fertilized seedlings. Increased NH(4)(+) availability significantly altered the distribution of fixed soil DIC among tissue types and increased the amount of fixed soil DIC in ectomycorrhizal roots by 130% compared with unfertilized seedlings. Increased NH(4)(+) availability did not increase fixation of soil DIC in nonmycorrhizal roots, suggesting that NH(4)(+) assimilation may be concentrated in ectomycorrhizal fungal tissues, reflecting greater anaplerotic demands. Soil DIC is likely to contribute only a small amount of C to forest trees, but it may be important in C fixation processes of specific tissues, such as newly formed stems and fine roots, and ectomycorrhizal roots assimilating NH(4)(+).