Photosynthate Transfer Research Articles

Iron Nanostructure Primes Arbuscular Mycorrhizal Fungi Symbiosis Tightly Connecting Maize Leaf Photosynthesis via a Nanofilm Effect.

It is crucial to clarify how the iron nanostructure activates plant growth, particularly in combination with arbuscular mycorrhizal fungi (AMF). We first identified 1.0 g·kg-1 of nanoscale zerovalent iron (nZVI) as appropriate dosage to maximize maize growth by 12.7-19.7% in non-AMF and 18.9-26.4% in AMF, respectively. Yet, excessive nZVI at 2.0 g·kg-1 exerted inhibitory effects while FeSO4 showed slight effects (p > 0.05). Under an appropriate dose, a nano core-shell structure was formed and the transfer and diffusion of electrons between PS II and PS I were facilitated, significantly promoting the reduction of ferricyanide and NADP (p < 0.05). SEM images showed that excessive nZVI particles can form stacked layers on the surface of roots and hyphae, inhibiting water and nutrient uptake. TEM observations showed that excessive nanoparticles can penetrate into root cortical cells, disrupt cellular homeostasis, and substantially elevate Fe content in roots (p < 0.05). This exacerbated membrane lipid peroxidation and osmotic regulation, accordingly restricting photosynthetic capacity and AMF colonization. Yet, appropriate nZVI can be adhered to a mycelium surface, forming a uniform nanofilm structure. The strength of the mycelium network was evidently enhanced, under an increased root colonization rate and an extramatrical hyphal length (p < 0.05). Enhanced mycorrhizal infection was tightly associated with higher gas exchange and Rubisco and Rubisco enzyme activities. This enabled more photosynthetic carbon to input into AMF symbiont. There existed a positive feedback loop connecting downward transfer of photosynthate and upward transport of water/nutrients. FeSO4 only slightly affected mycorrhizal development. Thus, it was the Fe nanostructure but not its inorganic salt state that primed AMF symbionts for better growth.

Photosynthate transfer from an autotrophic orchid to conspecific heterotrophic protocorms through a common mycorrhizal network.

The minute 'dust seeds' of some terrestrial orchids preferentially germinate and develop as mycoheterotrophic protocorms near conspecific adult plants. Here we test the hypothesis that mycorrhizal mycelial connections provide a direct pathway for transfer of recent photosynthate from conspecific green orchids to achlorophyllous protocorms. Mycelial networks of Ceratobasidium cornigerum connecting green Dactylorhiza fuchsii plants with developing achlorophyllous protocorms of the same species were established on oatmeal or water agar before the shoots of green plants were exposed to 14CO2. After incubation for 48 h, the pattern of distribution of fixed carbon was visualised in intact entire autotrophic/protocorm systems using digital autoradiography and quantified in protocorms by liquid scintillation counting. Both methods of analysis revealed accumulation of 14C above background levels in protocorms, confirming that autotrophic plants supply carbon to juveniles via common mycorrhizal networks. Despite some accumulation of plant-fixed carbon in the fungal mycelium grown on oatmeal agar, a greater amount of carbon was transferred to protocorms growing on water agar, indicating that the polarity of transfer may be influenced by sink strength. We suggest this transfer pathway may contribute significantly to the pattern and processes determining localised orchid establishment in nature, and that 'parental nurture' via common mycelial networks may be involved in these processes.

Olive Performance under the Soil Application of Humic Acid and the Spraying of Titanium and Zinc Nanoparticles under Soil Salinity Stress

Salinity is a major social, economic, and environmental menace in climates with low rainfall and high evapotranspiration, and it influences plant growth and causes restriction to crop production in the world. Additionally, under salinity stress, numerous physiological processes such as photosynthesis, biomass accumulation, and photosynthate transfer are also harshly lessened, and it also limits the absorption of adequate water by plants and leads to a dimension in plant water status. Therefore, the current study was conducted to investigate the soil application of humic acid (HA) at 0, 0.5, 1 and 2 kg/tree alone or in combination with the foliar spraying of 0 mg ZnO2 + 0 mg TiO2, 200 mg ZnO2 + 60 mg TiO2 and/or 300 mg ZnO2 + 80 mg TiO2 through the two successive seasons 2022 and 2023. The results demonstrated that the use of HA alone or in combination with the spraying of TiO2 and ZnO2 greatly improved the leaf chlorophyll, flower number, fruit set percentages, fruit yields in kg or in ton per hectare, fruit weight, fruit size, and fruit firmness. Additionally, the same used treatments greatly improved the fruit content from TSS and oil percentages and also the leaf mineral content from N, P and K, while they minimized the fruit drop percentage and fruit moisture content as compared to control. The most positive influence was observed with the soil implementation of 2 kg HA combined with 300 mg ZnO2 + 80 mg TiO2 in the two experimental seasons.

Polyethylene microplastics can attenuate soil carbon sequestration by reducing plant photosynthetic carbon assimilation and transfer: evidence from a 13C-labeling mesocosm study

Microplastics (MPs) are emerging environmental pollutants that potentially affect soil carbon (C) cycling. However, how MPs influence photosynthesized C flow from plants to soil remains unclear, creating large uncertainties in the projection of C dynamics in plant-soil-atmosphere systems. Herein, a 13C-labeling technique was applied to unravel the photosynthesized C fluxes from maize to soil under different levels of simulated polyethylene (PE) MPs pollution (0, 1% and 5%) in soil. Resultantly, the 1% and 5% PE MPs treatments significantly decreased the net fixed 13C in the maize–soil system. However, the effect of PE MPs on photosynthate–13C transfer in the maize–soil system was highly concentration–specific, that is, the addition of 1% PE MPs reduced the transfer of photosynthate–13C to the belowground part (−44.6%) and soil (−36.3%), whereas these effects were not observed for 5% PE MPs. PE MPs addition did not change soil moisture and bulk density. The addition of 1% PE MPs increased soil microbial biomass and decreased soil nitrogen (N) availability (inorganic and organic N). The treatment with 5% PE MPs decreased soil microbial community diversity, and altered the microbial community structure. Soil pH, organic N, and aboveground and belowground plant biomass mainly controlled the soil 13C content. These findings suggest that plant–soil systems polluted by PE MPs would sequester less photosynthesized C to belowground parts and soil through reducing plant photosynthetic carbon assimilation and transfer. Thus, potential changes in soil C stocks under MPs pollution should be considered for predicting global C dynamics and sustainable agricultural production.

Intensive grassland management disrupts below-ground multi-trophic resource transfer in response to drought

Modification of soil food webs by land management may alter the response of ecosystem processes to climate extremes, but empirical support is limited and the mechanisms involved remain unclear. Here we quantify how grassland management modifies the transfer of recent photosynthates and soil nitrogen through plants and soil food webs during a post-drought period in a controlled field experiment, using in situ 13C and 15N pulse-labelling in intensively and extensively managed fields. We show that intensive management decrease plant carbon (C) capture and its transfer through components of food webs and soil respiration compared to extensive management. We observe a legacy effect of drought on C transfer pathways mainly in intensively managed grasslands, by increasing plant C assimilation and 13C released as soil CO2 efflux but decreasing its transfer to roots, bacteria and Collembola. Our work provides insight into the interactive effects of grassland management and drought on C transfer pathways, and highlights that capture and rapid transfer of photosynthates through multi-trophic networks are key for maintaining grassland resistance to drought.

Large contribution of recent photosynthate to soil respiration in tropical dipterocarp forest revealed by girdling

Abstract Tropical forests are the most productive terrestrial ecosystems, fixing over 40 Pg of carbon from the atmosphere each year. A substantial portion of this carbon is allocated below‐ground to roots and root‐associated micro‐organisms. However, there have been very few empirical studies on the dynamics of this below ground transfer, especially in tropical forests where carbon allocation processes are influenced by high plant species diversity. We used a whole‐stand girdling experiment to halt the below‐ground transfer of recent photosynthates in a lowland tropical forest in Borneo. By girdling 209 large trees in a 0.48 ha plot, we determined: (a) the contribution of recent photosynthate to root‐rhizosphere respiration and; (b) the relationships among the disruption of this below‐ground carbon supply, tree species composition and mortality. Mortality of the 209 trees was 62% after 370 days, with large variation among species and particularly high mortality within the Dipterocarpaceae (99%) and Fagaceae (100%) families. We also observed a higher risk of mortality following girdling for species with lower wood density. Soil CO2 emissions declined markedly (36 ± 5%) over ~50 days following girdling in three of six monitored subplots. In the other three subplots there was either a marginal decline or no response of soil CO2 emissions to girdling. The decrease in soil CO2 efflux was larger in subplots with dominance of Dipterocarpaceae. Synthesis. Our results indicate high spatial variation in the coupling of below‐ground carbon allocation and root‐rhizosphere respiration in this tropical forest, with a closer coupling in forest dominated by Dipterocarpaceae. Our findings highlight the implications of tree species composition of tropical forests in affecting the dynamics of below‐ground carbon transfer and its release to the atmosphere.

Heterotrophic Carbon Fixation in a Salamander-Alga Symbiosis.

The unique symbiosis between a vertebrate salamander, Ambystoma maculatum, and unicellular green alga, Oophila amblystomatis, involves multiple modes of interaction. These include an ectosymbiotic interaction where the alga colonizes the egg capsule, and an intracellular interaction where the alga enters tissues and cells of the salamander. One common interaction in mutualist photosymbioses is the transfer of photosynthate from the algal symbiont to the host animal. In the A. maculatum–O. amblystomatis interaction, there is conflicting evidence regarding whether the algae in the egg capsule transfer chemical energy captured during photosynthesis to the developing salamander embryo. In experiments where we took care to separate the carbon fixation contributions of the salamander embryo and algal symbionts, we show that inorganic carbon fixed by A. maculatum embryos reaches 2% of the inorganic carbon fixed by O. amblystomatis algae within an egg capsule after 2 h in the light. After 2 h in the dark, inorganic carbon fixed by A. maculatum embryos is 800% of the carbon fixed by O. amblystomatis algae within an egg capsule. Using photosynthesis inhibitors, we show that A. maculatum embryos and O. amblystomatis algae compete for available inorganic carbon within the egg capsule environment. Our results confirm earlier studies suggesting a role of heterotrophic carbon fixation during vertebrate embryonic development. Our results also show that the considerable capacity of developing A. maculatum embryos for inorganic carbon fixation precludes our ability to distinguish any minor role of photosynthetically transferred carbon from algal symbionts to host salamanders using bicarbonate introduced to the egg system as a marker.

Clonal integration enhances performance of an invasive grass

While many clonal plants are highly successful invaders, the contribution of clonal integration (i.e. the translocation of resources among ramets) to invasion is often unknown. We used model simulations to ask if clonal integration would facilitate photosynthate translocation, if the performance of daughter ramets might be enhanced by clonal integration, and if shaded ramets benefited relatively more from transferred photosynthate. Then, to test if photosynthate translocation augmented performance of emerging daughter ramets for a globally invasive grass (Imperata cylindrica), we combined a 13CO2 pulse‐chase experiment with a greenhouse experiment manipulating light levels and rhizome attachment. We found that acropetal photosynthate transfer occurred between all sampled parent–daughter ramet pairs and that this resource sharing led to higher biomass and tiller production when rhizomes between parent and daughter ramets were intact. We also found that the benefits of integration to recipient clones outweighed the costs to donors, since there was no reduction in parent plant performance due to sharing. Additionally, our data analyses show that photosynthate transfer was likely of greater benefit in overcoming growth constraints in shade than in full sun (posterior probability ~96.5%), a result that is further supported by our numerical simulations from a basic growth model. Thus, resource sharing among clonal plants may be a critical but underappreciated trait of invasive species. More generally, photosynthate transfer is a probable mechanism that explains why clonal integration can be particularly beneficial in heterogeneous resource environments.

Local and Systemic Effects of Brassinosteroid Perception in Developing Phloem.

The plant vasculature is an essential adaptation to terrestrial growth. Its phloem component permits efficient transfer of photosynthates between source and sink organs but also transports signals that systemically coordinate physiology and development. Here, we provide evidence that developing phloem orchestrates cellular behavior of adjacent tissues in the growth apices of plants, the meristems. Arabidopsis thaliana plants that lack the three receptor kinases BRASSINOSTEROID INSENSITIVE 1 (BRI1), BRI1-LIKE 1 (BRL1), and BRL3 ("bri3" mutants) can no longer sense brassinosteroid phytohormones and display severe dwarfism as well as patterning and differentiation defects, including disturbed phloem development. We found that, despite the ubiquitous expression of brassinosteroid receptors in growing plant tissues, exclusive expression of the BRI1 receptor in developing phloem is sufficient to systemically correct cellular growth and patterning defects that underlie the bri3 phenotype. Although this effect is brassinosteroid-dependent, it cannot be reproduced with dominant versions of known downstream effectors of BRI1 signaling and therefore possibly involves a non-canonical signaling output. Interestingly, the rescue of bri3 by phloem-specific BRI1 expression is associated with antagonism toward phloem-specific CLAVATA3/EMBRYO SURROUNDING REGION-RELATED 45 (CLE45) peptide signaling in roots. Hyperactive CLE45 signaling causes phloem sieve element differentiation defects, and consistently, knockout of CLE45 perception in bri3 background restores proper phloem development. However, bri3 dwarfism is retained in such lines. Our results thus reveal local and systemic effects of brassinosteroid perception in the phloem: whereas it locally antagonizes CLE45 signaling to permit phloem differentiation, it systemically instructs plant organ formation via a phloem-derived, non-cell-autonomous signal.

Drought decreases incorporation of recent plant photosynthate into soil food webs regardless of their trophic complexity.

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO2 efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with 13C‐CO2 to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of 13C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, 13C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO2 efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment.

Rapid Transfer of Plant Photosynthates to Soil Bacteria via Ectomycorrhizal Hyphae and Its Interaction With Nitrogen Availability.

Plant roots release recent photosynthates into the rhizosphere, accelerating decomposition of organic matter by saprotrophic soil microbes (“rhizosphere priming effect”) which consequently increases nutrient availability for plants. However, about 90% of all higher plant species are mycorrhizal, transferring a significant fraction of their photosynthates directly to their fungal partners. Whether mycorrhizal fungi pass on plant-derived carbon (C) to bacteria in root-distant soil areas, i.e., incite a “hyphosphere priming effect,” is not known. Experimental evidence for C transfer from mycorrhizal hyphae to soil bacteria is limited, especially for ectomycorrhizal systems. As ectomycorrhizal fungi possess enzymatic capabilities to degrade organic matter themselves, it remains unclear whether they cooperate with soil bacteria by providing photosynthates, or compete for available nutrients. To investigate a possible C transfer from ectomycorrhizal hyphae to soil bacteria, and its response to changing nutrient availability, we planted young beech trees (Fagus sylvatica) into “split-root” boxes, dividing their root systems into two disconnected soil compartments. Each of these compartments was separated from a litter compartment by a mesh penetrable for fungal hyphae, but not for roots. Plants were exposed to a 13C-CO2-labeled atmosphere, while 15N-labeled ammonium and amino acids were added to one side of the split-root system. We found a rapid transfer of recent photosynthates via ectomycorrhizal hyphae to bacteria in root-distant soil areas. Fungal and bacterial phospholipid fatty acid (PLFA) biomarkers were significantly enriched in hyphae-exclusive compartments 24 h after 13C-CO2-labeling. Isotope imaging with nanometer-scale secondary ion mass spectrometry (NanoSIMS) allowed for the first time in situ visualization of plant-derived C and N taken up by an extraradical fungal hypha, and in microbial cells thriving on hyphal surfaces. When N was added to the litter compartments, bacterial biomass, and the amount of incorporated 13C strongly declined. Interestingly, this effect was also observed in adjacent soil compartments where added N was only available for bacteria through hyphal transport, indicating that ectomycorrhizal fungi were acting on soil bacteria. Together, our results demonstrate that (i) ectomycorrhizal hyphae rapidly transfer plant-derived C to bacterial communities in root-distant areas, and (ii) this transfer promptly responds to changing soil nutrient conditions.

Exudation rates and δ13C signatures of tree root soluble organic carbon in a riparian forest

Tree root exudation (TRE) of water soluble organic carbon (WSOC) is an important but under-assessed component of net primary production, and is thought to strongly influence rhizosphere biogeochemistry. Riparian systems in particular are often viewed as biogeochemical hot spots fueled partially by root exudate WSOC. However, TRE rates have not been previously reported for these systems. The δ13C signatures of exudates may provide important insights into plant physiology and inform isotope-based methods to identify sources of soil CO2 fluxes, but this information is also generally lacking. In the present study, root exudate WSOC was collected in situ to assess both net exudation rates and exudate δ13C values in a temperate riparian forest. Net TRE rates were found to be most strongly related to a combination of tree species, root characteristics and net ecosystem exchange (Adj. R2 = 0.73; p < 0.001). In contrast, exudate δ13C values were correlated to time-lagged vapor pressure deficit (Adj. R2 = 0.21; p < 0.05) and air temperature (Adj. R2 = 0.43; p < 0.05), suggesting a rapid transfer of photosynthate from the canopy to the rhizosphere. Extrapolation of mean net TRE rates (13 µmol C g root−1 day−1) from a root mass basis to the entire sampling area suggests that TRE may account for as much as 3% of net annual C uptake and represents an important input of organic matter to riparian soils. Our findings of predictable TRE rates and exudate δ13C values in the present study suggest that future studies examining δ13C values of different plant components, soil organic matter and respired soil CO2 would benefit by accounting for the impact of root exudates.

Response of Summer Maize Photosynthate Accumulation and Distribution to Shading Stress Assessed by Using 13CO2 Stable Isotope Tracer in the Field.

Maize is one of the most important crops globally that provides food, feed, and bioenergy. However, shading stress threatens maize production. In this study, we investigated the effects of shading on photosynthate accumulation and distribution of summer maize in the field. Zhengdan958 (ZD958) and Denghai 605 (DH605) were used as experimental materials in a field experiment running from 2013 to 2015. Shading treatments were applied over different growth stages: from the tassel stage (VT) to physiological maturity (R6) (S1), from the six-leaf stage (V6) to VT (S2), and from emergence stage (VE) to R6 (S3). The effects of shading on plant photosynthesis, photosynthate accumulation and distribution, and yield were evaluated in comparison to ambient sunlight. Shading significantly decreased the leaf area, SPAD value, net photosynthetic rate, dry matter accumulation, and grain yield. During the 3-year experimental period, grain yields of ZD958 and DH605 were reduced by 83.4%, 34.2%, 53.1% and 79.3%, 24.2%, 57.6% as compared to the CK by treatments S3, S2, and S1, respectively. 13CO2 stable isotope tracing revealed that shading differentially affected the photosynthate transfer rate in different stages; photosynthates were transferred from top to bottom plant parts, in the order control > S2 > S1 > S3. We conclude that shading clearly disrupted photosynthate metabolism, and reduced the photosynthate accumulation in the grain, resulting in a yield reduction.

Transcriptomic Analysis of Thermally Stressed Symbiodinium Reveals Differential Expression of Stress and Metabolism Genes.

Endosymbioses between dinoflagellate algae (Symbiodinium sp.) and scleractinian coral species form the foundation of coral reef ecosystems. The coral symbiosis is highly susceptible to elevated temperatures, resulting in coral bleaching, where the algal symbiont is released from host cells. This experiment aimed to determine the transcriptional changes in cultured Symbiodinium, to better understand the response of cellular mechanisms under future temperature conditions. Cultures were exposed to elevated temperatures (average 31°C) or control conditions (24.5°C) for a period of 28 days. Whole transcriptome sequencing of Symbiodinium cells on days 4, 19, and 28 were used to identify differentially expressed genes under thermal stress. A large number of genes representing 37.01% of the transcriptome (∼23,654 unique genes, FDR < 0.05) with differential expression were detected at no less than one of the time points. Consistent with previous studies of Symbiodinium gene expression, fold changes across the transcriptome were low, with 92.49% differentially expressed genes at ≤2-fold change. The transcriptional response included differential expression of genes encoding stress response components such as the antioxidant network and molecular chaperones, cellular components such as core photosynthesis machinery, integral light-harvesting protein complexes and enzymes such as fatty acid desaturases. Differential expression of genes encoding glyoxylate cycle enzymes were also found, representing the first report of this in Symbiodinium. As photosynthate transfer from Symbiodinium to coral hosts provides up to 90% of a coral’s daily energy requirements, the implications of altered metabolic processes from exposure to thermal stress found in this study on coral-Symbiodinium associations are unknown and should be considered when assessing the stability of the symbiotic relationship under future climate conditions.

Functional Characterization of a Hexose Transporter from Root Endophyte Piriformospora indica.

Understanding the mechanism of photosynthate transfer at symbiotic interface by fungal monosaccharide transporter is of substantial importance. The carbohydrate uptake at the apoplast by the fungus is facilitated by PiHXT5 hexose transporter in root endophytic fungus Piriformospora indica. The putative PiHXT5 belongs to MFS superfamily with 12 predicted transmembrane helices. It possess sugar transporter PFAM motif (PF0083) and MFS superfamily domain (PS50850). It contains the signature tags related to glucose transporter GLUT1 of human erythrocyte. PiHXT5 is regulated in response to mutualism as well as glucose concentration. We have functionally characterized PiHXT5 by complementation of hxt-null mutant of Saccharomyces cerevisiae EBY.VW4000. It is involved in transport of multiple sugars ranging from D-glucose, D-fructose, D-xylose, D-mannose, D-galactose with decreasing affinity. The uncoupling experiments indicate that it functions as H+/glucose co-transporter. Further, pH dependence analysis suggests that it functions maximum between pH 5 and 6. The expression of PiHXT5 is dependent on glucose concentration and was found to be expressed at low glucose levels (1 mM) which indicate its role as a high affinity glucose transporter. Our study on this sugar transporter will help in better understanding of carbon metabolism and flow in this agro-friendly fungus.

Molecular assessment of the effect of light and heterotrophy in the scleractinian coral Stylophora pistillata.

Corals acquire nutrients via the transfer of photosynthates by their endosymbionts (autotrophy), or via zooplankton predation by the animal (heterotrophy). During stress events, corals lose their endosymbionts, and undergo starvation, unless they increase their heterotrophic capacities. Molecular mechanisms by which heterotrophy sustains metabolism in stressed corals remain elusive. Here for the first time, to the best of our knowledge, we identified specific genes expressed in heterotrophically fed and unfed colonies of the scleractinian coral Stylophora pistillata, maintained under normal and light-stress conditions. Physiological parameters and gene expression profiling demonstrated that fed corals better resisted stress than unfed ones by exhibiting less oxidative damage and protein degradation. Processes affected in light-stressed unfed corals (HLU), were related to energy and metabolite supply, carbohydrate biosynthesis, ion and nutrient transport, oxidative stress, Ca(2+) homeostasis, metabolism and calcification (carbonic anhydrases, calcium-transporting ATPase, bone morphogenetic proteins). Two genes (cp2u1 and cp1a2), which belong to the cytochrome P450 superfamily, were also upregulated 249 and 10 times, respectively, in HLU corals. In contrast, few of these processes were affected in light-stressed fed corals (HLF) because feeding supplied antioxidants and energetic molecules, which help repair oxidative damage. Altogether, these results show that heterotrophy helps prevent the cascade of metabolic problems downstream of oxidative stress.

Pathway and sink activity for photosynthate translocation in Pisolithus extraradical mycelium of ectomycorrhizal Pinus thunbergii seedlings.

The purpose of this study was to identify the pathway and sink activity of photosynthate translocation in the extraradical mycelium (ERM) of a Pisolithus isolate. We labelled ectomycorrhizal (ECM) Pinus thunbergii seedlings with (14)CO2 and followed (14)C distribution within the ERM by autoradiography. (14)C photosynthate translocation in the ERM resulted in (14)C distribution in rhizomorphs throughout the ERM, with (14)C accumulation at the front. When most radial mycelial connections between ECM root tips and the ERM front were cut, the whole allocation of (14)C photosynthates to the ERM was reduced. However, the overall pattern of (14)C distribution in the ERM was maintained even in regions immediately above and below the cut, with no local (14)C depletion or accumulation. We inferred from this result that every portion in the ERM has a significant sink activity and a definite sink capacity for photosynthates and that photosynthates detour the cut and reach throughout the ERM by translocation in every direction. Next, we prepared paired ECM seedlings, ERMs of which had been connected with each other by hyphal fusion, alongside, labelled the left seedling with (14)CO2, and shaded none, one or both of them. (14)C photosynthates were acropetally and basipetally translocated from the left ERM to ECM root tips of the right seedling through rhizomorphs in the left and right ERMs, respectively. With the left seedling illuminated, (14)C translocation from the left to the right ERM increased by shading the right seedling. This result suggests that reduced photosynthate transfer from the host to its ERM increased sink activity of the ERM.

Stable isotopes in tree rings: towards a mechanistic understanding of isotope fractionation and mixing processes from the leaves to the wood.

The mechanistic understanding of isotope fractionation processes is increasing but we still lack detailed knowledge of the processes that determine the isotopic composition of the tree-ring archive over the long term. Especially with regard to the path from leaf photosynthate production to wood formation, post-assimilation fractionations/processes might cause at least a partial decoupling between the leaf isotope signals that record processes such as stomatal conductance, transpiration and photosynthesis, and the wood or cellulose signals that are stored in the paleophysiological record. In this review, we start from the rather well understood processes at the leaf level such as photosynthetic carbon isotope fractionation, leaf water evaporative isotope enrichment and the issue of the isotopic composition of inorganic sources (CO2 and H2O), though we focus on the less explored 'downstream' processes related to metabolism and transport. We further summarize the roles of cellulose and lignin as important chemical constituents of wood, and the processes that determine the transfer of photosynthate (sucrose) and associated isotopic signals to wood production. We cover the broad topics of post-carboxylation carbon isotope fractionation and of the exchange of organic oxygen with water within the tree. In two case studies, we assess the transfer of carbon and oxygen isotopic signals from leaves to tree rings. Finally we address the issue of different temporal scales and link isotope fractionation at the shorter time scale for processes in the leaf to the isotopic ratio as recorded across longer time scales of the tree-ring archive.

Origin and Evolution of Plastids and Photosynthesis in Eukaryotes

Recent progress in understanding the origins of plastids from endosymbiotic cyanobacteria is reviewed. Establishing when during geological time the endosymbiosis occurred remains elusive, but progress has been made in defining the cyanobacterial lineage most closely related to plastids, and some mechanistic insight into the possible existence of cryptic endosymbioses perhaps involving Chlamydia-like infections of the host have also been presented. The phylogenetic affinities of the host remain obscure. The existence of a second lineage of primary plastids in euglyphid amoebae has now been confirmed, but the quasipermanent acquisition of plastids by animals has been shown to be more ephemeral than initially suspected. A new understanding of how plastids have been integrated into their hosts by transfer of photosynthate, by endosymbiotic gene transfer and repatriation of gene products back to the endosymbiont, and by regulation of endosymbiont division is presented in context.

Sex-specific interaction between arbuscular mycorrhizal and dark septate fungi in the dioecious plantAntennaria dioica(Asteraceae)

Male and female plants of dioecious species often differ in their resource demands and this has been linked to secondary sexual dimorphism, including sex-specific interactions with other organisms such as herbivores and pollinators. However, little is known about the interaction between dioecious plants and fungal root endophytes. Plants may be simultaneously colonised by arbuscular mycorrhizal (AM) and dark septate (DS) fungi. While it is well established that AM mutualism involves reciprocal transfer of photosynthates and mineral nutrients between roots of host plants and these fungi, the role of DS fungi remains controversial. Here, we report the temporal and spatial variation in AM and DS fungi in female, male and non-reproductive Antennaria dioica plants in three natural populations in Finland during flowering and after seed production. Females had higher colonisation by AM fungi, but lower colonisation by DS fungi than male and non-reproductive plants. The higher AM colonisation was observed during flowering, and this difference varied among populations. Our results suggest that females and males of A.dioica interact with AM and DS fungi differently and that this relationship is dependent on soil fertility.