Photosynthetic Layer Research Articles

Influence of land-terminating glacier on primary production in the high Arctic fjord (Blagopoluchiya Bay, Novaya Zemlya archipelago, Kara Sea)

Glaciers retreat linked with rapid Arctic warming changes bioproductivity in the bays and coastal areas of the Arctic Ocean (AO). The impact of different types of glaciers shrinking on water column primary production (IPP) is widely discussed. This work studied the influence of the land-terminating glacier on IPP and chlorophyll a concentration (Chlph) in Blagopoluchiya Bay (Novaya Zemlya archipelago). It is known that meltwater runoff from land-terminating glaciers enriches coastal waters with inorganic suspended matter and nutrients. The aim of the present study was to establish the main environmental factor determining IPP in the bay influenced by the glacier. During summer and autumn, IPP in the bay was low. The medians of IPP were 69 and 26 mgC m−2 day−1, respectively, which was 1.7- and 1.9-fold lower than in open waters of the Kara Sea in the vicinity of the bay. Generally, IPP increased from the inner part, close to the glacier, to the outer part of the bay. In summer, IPP and Chlph depend on both nutrient concentrations in the photosynthetic layer and the parameters of water transparency. On the contrary, in autumn, IPP and Chlph depend only on the optical properties of the subsurface layer. It can be concluded that low IPP in Blagopoluchiya Bay is determined mainly by unfavorable conditions of underwater PAR formed by meltwater discharge from the glacier. Thus, the fast melting of the land-terminating glaciers linked with climate warming can lead to a decrease in the biological productivity of the AO bays.

Factors behind the threshold‐like changes in lake ecosystems along a water colour gradient: The effects of dissolved organic carbon and iron on euphotic depth, mixing depth and phytoplankton biomass

Abstract The effects of water colour, dissolved organic carbon (DOC), iron (Fe) and fetch on the mixing depth (Zmix), euphotic depth (Zeu) and phytoplankton biomass of lakes were studied to clarify the factors behind changes in lake productivity along a DOC gradient. The study was conducted by field sampling 102 lakes in Finland. The lakes were situated mainly in forested areas. At DOC concentrations below 15 mg/L, light attenuation was explained both by Fe and DOC, but at higher DOC concentrations the effect of Fe faded. For the absorption of different wavelengths of photosynthetic light, both DOC and Fe had a significant effect up to 475 nm, but for longer wavelengths only the effect of DOC was significant. Both Zmix and Zeu decreased logarithmically with increasing DOC and water colour. Water colour was determined by DOC and Fe. Zmix increased along with increasing fetch. Zmix exceeded Zeu when DOC concentration was >14.4 mg/L and water colour >106 mg Pt/L (platinum as chloroplatinate ion 3). Above the 14.4 mg/L DOC threshold, the illuminated part of the mixed layer decreased logarithmically with increasing water colour, leaving an increasing part of the mixed epilimnion below the photosynthetic layer. Chlorophyll‐a concentration showed a unimodal relationship with DOC, with the highest biomass occurring at DOC concentrations of 15.7–15.9 mg/L. The results suggested that above DOC concentration 14 mg/L and water colour 106 mg Pt/L, both quality and quantity of light in the epilimnion restrict production by phytoplankton. This can explain the observed threshold‐like changes in lake productivity along a DOC gradient.

Development of suberized barrier is critical for ion partitioning between senescent and non-senescent tissues in a succulent halophyte Sarcocornia quinqueflora

Senescence of plant tissues is a physiologically synchronized process that enables an evergreen or a perennial plant to retrieve, recycle and remobilize nutrients from elder to younger tissues or upcoming seeds. The succulent perennial halophyte S. quinqueflora utilizes this process to discard excess salt being accumulated in outer tissues of their leafy stems. The exact mechanism for salt shedding in this plant, however, remains elusive. In this work we show that the plant develops two distinct types of tissues - an endodermis-like layer (suberized layer, ED), and an additional internal photosynthetic layer (IP) - to enable this process. Their potential roles toward salt-coping strategy were investigated in this study. We show that elevated salinity leads to an accelerated development of the ED, and that its development strongly affected ion partitioning between outer (senescent) and inner (non-senescent) tissues. A positive correlation between the ratio of ED to a bead diameter and the outer to inner concentration of Na+ was observed. These ratios were highest in older (basipetally-located) beads and progressively decreased towards the tip. Furthermore, the Na+/ K+ ratio in inner tissues of bottom beads at highest salinity treatments (800 and 1000 mM NaCl) that showed clear senescence symptoms was ∼1.0, indicative of complete separation of the outer and inner tissues at late developmental stage due to the fully suberized ED multilayer. A dual-sources: dual-sinks model explaining the role of the IP layer in plant adaptation to salinity is presented.

RNA-Seq analysis reveals the growth and photosynthetic responses of rapeseed (Brassica napus L.) under red and blue LEDs with supplemental yellow, green, or white light

Compound light is required for plant growth and development, but the response mechanisms of plants are undercharacterized and not fully understood. The present study was undertaken to evaluate the effect of supplemental light (green light, G; white light, W; yellow light, Y) added to red–blue light (RB) and sole W on the growth and photosynthesis of rapeseed seedlings. The results revealed that supplemental G/W improved the growth and photosynthesis of seedlings, but supplemental Y significantly reduced the photosynthetic rate and palisade tissue layer. Sole W caused similar responses in terms of growth, leaf development, oxidative damage, and antioxidant capability as supplemental Y. In total, 449, 367, 813, and 751 differentially expressed genes (DEGs) were identified under supplemental G, Y, and W and sole W, respectively, compared to RB. The DEGs under different lights were closely associated with pathways such as light stimulus and high-light response, root growth, leaf development, photosynthesis, photosynthesis-antenna proteins, carbohydrate synthesis and degradation, secondary metabolism, plant hormones, and antioxidant capacity, which contributed to the distinct growth and photosynthesis under different treatments. Our results suggest that Y is more likely substituted by other wavelengths to achieve certain effects similar to those of supplemental Y, while G has a more distinctive effect on rapeseed. Taken together, supplementation RB with G/W promotes the growth of rapeseed seedlings in a controlled environment.

Differential gene expression associated with fungal trophic shifts along the senescence gradient of the moss Dicranum scoparium.

Bryophytes harbour microbiomes, including diverse communities of fungi. The molecular mechanisms by which perennial mosses interact with these fungal partners along their senescence gradients are unknown, yet this is an ideal system to study variation in gene expression associated with trophic state transitions. We investigated differentially expressed genes of fungal communities and their host Dicranum scoparium across its naturally occurring senescence gradient using a metatranscriptomic approach. Higher activity of fungal nutrient-related (carbon, nitrogen, phosphorus and sulfur) transporters and Carbohydrate-Active enZyme (CAZy) genes was detected toward the bottom, partially decomposed, layer of the moss. The most prominent variation in the expression levels of fungal nutrient transporters was from inorganic nitrogen-related transporters, whereas the breakdown of organonitrogens was detected as the most enriched gene ontology term for the host D. scoparium, for those transcripts having higher expression in the partially decomposed layer. The abundance of bacterial rRNA transcripts suggested that more living members of Cyanobacteria are associated with the photosynthetic layer of D. scoparium, while members of Rhizobiales are detected throughout the gametophytes. Plant genes for specific fungal-plant communication, including defense responses, were differentially expressed, suggesting that different genetic pathways are involved in plant-microbe crosstalk in photosynthetic tissues compared to partially decomposed tissues.

Reconstruction of phytoplankton productivity and community structure in the South Yellow Sea

The sedimentary environment and ecological system in the South Yellow Sea (SYS) changed dramatically due to sea level change caused by glacial-interglacial cycles. The authors report the use of marine biomarkers (brassicasterol, dinosterol and C₃₇ alkenones) and terrigenous biomarkers (C₂₈+C₃₀+C₃₂ <i>n</i>-alkanols) in core DLC70-3 from the SYS to reconstruct the variation in the phytoplankton productivity and community structure and possible mechanisms during the middle Pleistocene. The results show that the primary productivity and that of single algae presented a consistent trend for the whole core during the middle Pleistocene, which was high during interglacial periods and low during glacial periods, with the highest being in marine isotope stage (MIS) 5–9 and MIS 19–21. The main reason is that the Yellow Sea Warm Current (YSWC) carried much of high temperature, high salinity water into the SYS, causing upwelling and vertical mixing and stirring, which increased the nutrient supply in the photosynthetic layer. The phytoplankton community structure mainly showed an increase in the relative content of haptophytes in MIS 5–9 and MIS 19–21, while the relative content of diatoms and dinoflagellates decreased; there was no evidence for a haptophyte content in other stages. The results reveal a shift from a coccolitho-phorid-dominated community during MIS 5–9 and MIS 19–21 to a diatom-dominated community during the other stages, mainly as a result of surface salinity variation, attributed to the invasion of the YSWC during high sea level periods.

Phytophthora palmivora establishes tissue-specific intracellular infection structures in the earliest divergent land plant lineage

The expansion of plants onto land was a formative event that brought forth profound changes to the earth's geochemistry and biota. Filamentous eukaryotic microbes developed the ability to colonize plant tissues early during the evolution of land plants, as demonstrated by intimate, symbiosis-like associations in >400 million-year-old fossils. However, the degree to which filamentous microbes establish pathogenic interactions with early divergent land plants is unclear. Here, we demonstrate that the broad host-range oomycete pathogen Phytophthora palmivora colonizes liverworts, the earliest divergent land plant lineage. We show that P. palmivora establishes a complex tissue-specific interaction with Marchantia polymorpha, where it completes a full infection cycle within air chambers of the dorsal photosynthetic layer. Remarkably, P. palmivora invaginates M. polymorpha cells with haustoria-like structures that accumulate host cellular trafficking machinery and the membrane syntaxin MpSYP13B, but not the related MpSYP13A. Our results indicate that the intracellular accommodation of filamentous microbes is an ancient plant trait that is successfully exploited by pathogens like P. palmivora.

Sediment Flux of Particulate Organic Phosphorus in the Open Black Sea

The interannual variation of the monthly average (weighted average) concentrations of particulate organic phosphorus (PPOM) in the photosynthetic layer, oxycline, redox zone, and H2S zone in the open Black Sea is estimated based on long-term observation data. The suspension sedimentation rates from the studied layers are assessed using model calculations and published data. The annual variation of PPOM sediment fluxes from the photosynthetic layer, oxycline, redox zone, and upper H2S zone to the anaerobic zone of the sea and the correspondingly annual average values are estimated for the first time. A regular decrease in the PPOM annual average flux with depth in the upper active layer is demonstrated. A correlation between the annual average values of PPOM sediment flux from the photosynthetic layer and ascending phosphate flux to this layer is shown, which suggests their balance in the open sea. The results are discussed in terms of the phosphorus biogeochemical cycle and the concept of new and regenerative primary production in the open Black Sea.

Characterisation of estuarine intertidal macroalgae by laser-induced fluorescence

The article reports the application of laser-induced fluorescence (LIF) for the assessment of macroalgae communities of estuarine intertidal areas. The method was applied for the characterisation of fifteen intertidal macroalgae species of the Tagus estuary, Portugal, and adjacent coastal area. Three bands characterised the LIF spectra of red macroalgae with emission maxima in the ranges 577–583 nm, 621–642 nm and 705–731 nm. Green and brown macroalgae showed one emission maximum in the red region (687–690 nm) and/or one in the far-red region (726–732 nm). Characteristics of LIF emission spectra were determined by differences in the main fluorescing pigments: phycoerythrin, phycocyanin and chlorophyll a (Chl a). In the green and brown macroalgae groups, the relative significance of the two emission maxima seems to be related to the thickness of the photosynthetic layer. In thick macroalgae, like Codium tomentosum or Fucus vesiculosus, the contribution of the far-red emission fluorescence peak was more significant, most probably due to re-absorption of the emitted red Chl a fluorescence within the dense photosynthetic layer. Similarly, an increase in the number of layers of the thin-blade green macroalgae Ulva rigida caused a shift to longer wavelengths of the red emission maximum and the development of a fluorescence peak at the far-red region. Water loss from Ulva's algal tissue also led to a decrease in the red/far-red Chl fluorescence ratio (F685/F735), indicating an increase in the density of chloroplasts in the shrinking macroalgal tissue during low tide exposure.

Evaluation of the influence of abiotic and biotic factors on primary production in the Kara Sea in autumn

A regression analysis of the parameters of primary production versus environmental factors was performed on the basis of data of three complex expeditions performed in the Kara Sea in September to October 1993, 2007, and 2011. The analysis of the dependence of the depth-integrated primary production (PPint) on the value of surface chlorophyll a (Chl0) and assimilation activity (ANm) showed that only 12% of the variability of the integrated PPint was determined by the variability of Chl0, whereas the correlation between PPint and ANm was strong (R2 = 0.635). Thus, in autumn, PPint values in the Kara Sea depended primarily on the activity of phytoplankton assimilation. At the end of the vegetative season, high (close to or above 1 mg/m3) Chl0 values did not reflect phytoplankton production within the entire photosynthetic layer, where organic matter was synthesized at a low rate. In turn, PPint and ANm depended primarily on the intensity of insolation and was weakly related to the content of dissolved forms of nitrogen and phosphorus. In autumn, at the end of the vegetative season, insolation apparently is the main factor in the determination of the formation of primary production in the Kara Sea.

Disruption of photoautotrophic intertidal mats by filamentous fungi.

Ring-like structures, 2.0-4.8 cm in diameter, observed in photosynthetic microbial mats on the Wadden Sea island Schiermonnikoog (the Netherlands) showed to be the result of the fungus Emericellopsis sp. degrading the photoautotrophic top layer of the mat. The mats were predominantly composed of cyanobacteria and diatoms, with large densities of bacteria and viruses both in the top photosynthetic layer and in the underlying sediment. The fungal attack cleared the photosynthetic layer; however, no significant effect of the fungal lysis on the bacterial and viral abundances could be detected. Fungal-mediated degradation of the major photoautotrophs could be reproduced by inoculation of non-infected mat with isolated Emericellopsis sp., and with an infected ring sector. Diatoms were the first re-colonizers followed closely by cyanobacteria that after about 5 days dominated the space. The study demonstrated that the fungus Emericellopsis sp. efficiently degraded a photoautotrophic microbial mat, with potential implications for mat community composition, spatial structure and productivity.

Microbial characterization of microbial ecosystems associated to evaporites domes of gypsum in Salar de Llamara in Atacama desert.

The Central Andes in northern Chile contains a large number of closed basins whose central depression is occupied by saline lakes and salt crusts (salars). One of these basins is Salar de Llamara (850 m a.s.l.), where large domed structures of seemingly evaporitic origin forming domes can be found. In this work, we performed a detailed microbial characterization of these domes. Mineralogical studies revealed gypsum (CaSO(4)) as a major component. Microbial communities associated to these structures were analysed by 454 16S rDNA amplicon sequencing and compared between winter and summer seasons. Bacteroidetes Proteobacteria and Planctomycetes remained as the main phylogenetic groups, an increased diversity was found in winter. Comparison of the upper air-exposed part and the lower water-submerged part of the domes in both seasons showed little variation in the upper zone, showing a predominance of Chromatiales (Gammaproteobacteria), Rhodospirillales (Alphaproteobacteria), and Sphingobacteriales (Bacteroidetes). However, the submerged part showed marked differences between seasons, being dominated by Proteobacteria (Alpha and Gamma) and Verrucomicrobia in summer, but with more diverse phyla found in winter. Even though not abundant by sequence, Cyanobacteria were visually identified by scanning electron microscopy (SEM), which also revealed the presence of diatoms. Photosynthetic pigments were detected by high-performance liquid chromatography, being more diverse on the upper photosynthetic layer. Finally, the system was compared with other endoevaporite, mats microbialite and Stromatolites microbial ecosystems, showing higher similitude with evaporitic ecosystems from Atacama and Guerrero Negro. This environment is of special interest for extremophile studies because microbial life develops associated to minerals in the driest desert all over the world. Nevertheless, it is endangered by mining activity associated to copper and lithium extraction; thus, its environmental protection preservation is strongly encouraged.

Patterns of the Kara Sea primary production in autumn: Biotic and abiotic forcing of subsurface layer

Primary production and fundamental environmental factors were measured during September–October 1993, 2007 and 2011 in the Kara Sea. Relationships between the depth-integrated primary production (PPint), the surface chlorophyll a (Chl0) concentration and the maximum chlorophyll specific carbon rate within water column (Pbopt) had shown that only 12% of PPint variability were determined by Chl0 and there were strong correlations between PPint and Pbopt (R2=0.64). Thus, in the autumn PPint values were largely influenced by the phytoplankton assimilation activity. At the end of a vegetative season high (close or above 1mgm−3) values of Chl0 were not an index of phytoplankton productivity within photosynthetic layer where the organic matter synthesis rate was low which testifies the lack of correlation between Chl0 and PPint (R2=0.12) and between Chl0 and depth-integrated chlorophyll a (chl a) (R2=0.22). This conclusion is based on the low assimilation activity within water column and small thickness of the photosynthetic layer. The latter corresponds to low insolation and water transparency. The lack of correlation between surface and depth-integrated productivity parameters is the challenge for satellite PPint estimations. In turn PPint and Pbopt depended mainly on photosynthetically available radiation (PAR) and weakly related to the nitrogen and phosphorus concentrations. At the end of a vegetative season PAR level, apparently, should be considered as the main factor for primary production in the Kara Sea. Comparison between the integrated primary production from shipboard in September and the simulated evaluations of productivity (PP models) suggests that PP models overestimate the in situ PPint by a factor of 3–7 in the different Kara Sea regions. Improving of Kara Sea primary production estimations implies the development of regional satellite chl a algorithm and local primary production model considering specific features of PP in this Arctic Ocean region.

Difference of Characteristics of Photosynthesis, Matter Production and Translocation between Indica and Japonica Super Rice

This study was to reveal the difference of characteristics of dry matter accumulation and photosynthetic production between indica and japonica super rice. Field experiments were conducted by using five representative super hybrid indica combinations and five conventional japonica super rice varieties planted on a large scale in Jiangsu region with rice-wheat double cropping rotation. Dry matter accumulation, distribution and translocation, leaf area, leaf area duration(LAD), crop growth rate(CGR), net assimilation rate(NAR), seedling quality and leaf type were analyzed systematically. Results showed that, dry matter accumulation, LAD, CGR, and NAR at the early growth stage(from transplanting to jointing), leaf length, leaf base angle, angle between stem and leaf and drooping angle in top three leaves, leaf area decreasing per day after heading and harvest index of japonica rice were less than those of indica rice significantly, while dry matter accumulation, LAD, CGR, and NAR at the middle and later growth stages(from jointing to maturity), ratio of leaf area of productive tillers, ratio of leaf area of top three leaves, grain-leaf ratio(spikelets per cm2 leaf area, filled grains per cm2 leaf area and grain weight per cm2 leaf area), maximum leaf area index, total filling, grain yield, biological yield, maximum output and apparent output and their ratio were larger than those ofindica rice significantly. Although dry matter weight per stem of japonica rice at the main growth stages was lower than that of indica rice, the superiority of population quantity suggested that japonica rice would possess higher dry matter accumulation and leaf area. With the advancement of growth and development, the superiority of photosynthesis and matter production of japonica rice was increasing, with a higher dry matter accumulation more than that of indica rice 25 d after heading approximately. The growth potential of japonica rice remained strong at the late period of grain-filling(from milky stage to maturity), while stored matter kept in the stem and sheath exported appropriately at the early period of grain-filling(from heading to milky stage), ensuring the security of efficient photosynthetic layer and the enrichment of high accumulation yield sink. Therefore, balanced formation of high biological yield, stable-increasing-slow-decreasing tendency of leaf area, and the large amounts of high efficient photosynthate production from jointing to maturity are the important characteristics and causes for the efficient and sustainable output of photosynthetic systems, more grain-filling and high yield formation in japonica rice.

Variability of suspended organic matter in the northwestern Black Sea

The peculiarities in the distribution pattern of the concentration of suspended organic matter (SOM) components were studied for the long-term period of 1979–1995. We have analyzed their seasonal, annual, and interannual variability in the surface water layer and in the photosynthetic water layer in the northwestern Black Sea. Four areas pertaining to different concentrations of SOM components were defined according to the effect of riverine discharge and of open sea water masses, i.e. western, northern, central, and eastern areas. We have found an increase in the concentrations of SOM components in the summer-autumn period in the late 1980s and early 1990s, with the highest values observed in 1992. The concentration of suspended organic carbon was estimated for the concentration of Chlorophyll a in the surface water layer. These calculations are based on the regression relationship between simultaneous measurements of these two parameters in situ. It was found that the seasonal interannual variability in the concentrations of SOM components was affected by the volume of riverine discharge, its run to the sea, and climatic shifts.

Photosynthesis Assessment in Microphytobenthos Using Conventional and Imaging Pulse Amplitude Modulation Fluorometry

Imaging pulse amplitude modulated (Imaging-PAM) fluorometry is a breakthrough in the study of spatial heterogeneity of photosynthetic assemblages. However, Imaging and conventional PAM uses a different technology, making comparisons between these techniques doubtful. Thereby, photosynthetic processes were comparatively assessed using conventional (Junior PAM and PAM 101) and Imaging-PAM on intertidal microphytobenthos (MPB; mud and sand) and on cork oak leaves. Lower values of α (initial slope of the rETR, relative photosynthetic electron transport rate) vs E (incident photosynthetic active radiation) curve), ETR(max) (maximum relative ETR), E(k) (light saturation parameter) and F(v)/F(m) (maximum quantum efficiency of photosystem II of dark-adapted samples) were obtained using the Imaging-PAM. The level of discrepancy between conventional and Imaging-PAM systems was dependent on the type of sample, being more pronounced for MPB muddy sediments. This may be explained by differences in the depth integration of the fluorescence signal related to the thickness of the photosynthetic layer and in the light attenuation coefficients of downwelling irradiance. An additional relevant parameter is the taxonomic composition of the MPB, as cyanobacteria present in sandy sediments rendered different results with red and blue excitation light fluorometers. These findings emphasize the caution needed when interpreting chlorophyll fluorescence data of MPB communities.

Meridional asymmetric distribution of the primary production in the Atlantic Sector of the Southern Ocean in the austral spring and summer

The spatial distribution of the phytoplankton productivity was investigated in the Atlantic Sector of the Southern Ocean in the austral summer of 2009–2010 and the spring of 2010 on the basis of field measurements. In October–November, the average integrated primary production and the concentration of surface and integrated (photosynthetic layer) chlorophyll “a” in the Subantarctic waters of the Drake Passage exceeded the similar values along the Greenwich meridian by 3, 2, and 1.5 times, respectively. Similar primary production was observed in December–January for the water column in the eastern and western sectors of the studied area. The chlorophyll “a” concentration in the surface water layer was higher by 1.7 times for the Greenwich meridian area compared to the deep layers, but the concentrations of this pigment in the deeper layers did not differ. During the spring, the average primary production in the water column (the chlorophyll “a” concentration in the surface and in the photosynthetic layer) differed in the Drake Passage and along the Greenwich meridian by 2.3, 1.6, and 1.7 times, respectively. The opposite pattern was observed during the summer period, when the parameters listed above were higher for the Greenwich meridian area by 1.9, 2.5, and 1.7 times, respectively. Therefore, the western Antarctic areas in the spring are characterized by higher production than the eastern ones, and an opposite pattern is observed in the summer. The possible reasons for the meridional zonation of the chlorophyll “a” and the primary production are discussed in regard to different seasons.

Effects of temperature and rainfall variation on population structure and sexual dimorphism across the geographical range of a dioecious species

AbstractThe effects of climate (precipitation and temperature) on sexual dimorphism and population structure were analysed along a broad‐scale environmental gradient covering the distributional range of the endemic dioecious species Corema album, along the west coast of the Iberian Peninsula. We aimed to assess distribution constraints and sex‐related differences in demography and size associated with higher reproductive investment in females. Nine populations were chosen from across the geographic range of C. album and ten 10 × 10 m plots were established (10 m apart) along a 200‐m transect. All male, female and non‐reproductive shrubs were quantified within each plot and plant size, photosynthetic layer, height, sex ratio, population density and structure, and spatial segregation of sexes, under environmental conditions ranging from temperate to Mediterranean climate, were recorded and analysed. Increased aridity was related to lower population density and less structured populations, indicating an effect of higher temperature and lower precipitation on regeneration. Sexual dimorphism was influenced by climate, with size differences between sexes varying with aridity. However, demographic differences between sexes reflected in sex ratio deviations or the occurrence of spatial segregation were unrelated to any climatic variable, suggesting the existence of compensatory mechanisms that may counterbalance the higher reproductive effort of female plants. The results show the vulnerability of this endemic species to the increase in aridity expected in the southernmost limit of the biogeographical area due to global climate change, and demonstrate the importance of broad scale studies in the assessment of sexual dimorphism.

Cyanolichens can have both cyanobacteria and green algae in a common layer as major contributors to photosynthesis.

Cyanolichens are usually stated to be bipartite (mycobiont plus cyanobacterial photobiont). Analyses revealed green algal carbohydrates in supposedly cyanobacterial lichens (in the genera Pseudocyphellaria, Sticta and Peltigera). Investigations were carried out to determine if both cyanobacteria and green algae were present in these lichens and, if so, what were their roles. The types of photobiont present were determined by light and fluorescence microscopy. Small carbohydrates were analysed to detect the presence of green algal metabolites. Thalli were treated with selected strengths of Zn(2+) solutions that stop cyanobacterial but not green algal photosynthesis. CO(2) exchange was measured before and after treatment to determine the contribution of each photobiont to total thallus photosynthesis. Heterocyst frequencies were determined to clarify whether the cyanobacteria were modified for increased nitrogen fixation (high heterocyst frequencies) or were normal, vegetative cells. Several cyanobacterial lichens had green algae present in the photosynthetic layer of the thallus. The presence of the green algal transfer carbohydrate (ribitol) and the incomplete inhibition of thallus photosynthesis upon treatment with Zn(2+) solutions showed that both photobionts contributed to the photosynthesis of the lichen thallus. Low heterocyst frequencies showed that, despite the presence of adjacent green algae, the cyanobacteria were not altered to increase nitrogen fixation. These cyanobacterial lichens are a tripartite lichen symbiont combination in which the mycobiont has two primarily photosynthetic photobionts, 'co-primary photobionts', a cyanobacterium (dominant) and a green alga. This demonstrates high flexibility in photobiont choice by the mycobiont in the Peltigerales. Overall thallus appearance does not change whether one or two photobionts are present in the cyanobacterial thallus. This suggests that, if there is a photobiont effect on thallus structure, it is not specific to one or the other photobiont.

Peperomia leaf cell wall interface between the multiple hypodermis and crystal-containing photosynthetic layer displays unusual pit fields

Leaves of succulent Peperomia obtusifolia (Piperaceae), and its related species, contain a large multilayered hypodermis (epidermis) subtended by a very small single-layered photosynthetic palisade parenchyma, the latter containing spherical aggregates of crystals called druses. Each druse is in a central vacuole surrounded by chloroplasts. All hypodermal cell walls are thin, except for thick lowermost periclinal walls associated with the upper periclinal walls of the subtending palisade cells. These thick walls display 'quilted' impressions (mounds) formed by many subtending palisade cells. Conspicuous depressions occur in most mounds, and each depression contains what appear to be many plasmodesmata. These depressions are opposite similar regions in adjacent thin palisade periclinal walls, and they can be considered special pit fields that represent thin translucent regions ('windows' or 'skylights'). Druses in the vacuoles of palisade cells occur below these pit field regions and are surrounded by conspicuous cytoplasmic chloroplasts with massive grana oriented perpendicular to the crystals, probably providing for an efficient photosynthetic system under low-intensity light. Leaf clearings and fractures, light microscopy and crossed polarizers, general and histochemical staining, and transmission and scanning electron microscopy were used to examine these structures. Druses in the vacuoles of palisade cells occur below the thin pit field regions in the wall interface, suggesting an interesting physical relationship that could provide a pathway for light waves, filtered through the multiple hypodermis. The light waves pass into the palisade cells and are collected and dispersed by the druses to surrounding chloroplasts with large grana. These results imply an intriguing possible efficient photosynthetic adaptation for species growing in low-light environments, and provide an opportunity for future research on how evolution through environmental adaptation aids plants containing crystals associated with photosynthetic tissues to exist under low-light intensity and with other stresses.