The effect of samara wing presence on predation of Acer pseudoplatanus (Sapindaceae) seeds on the ground

The key selective pressure shaping the morphology of samaras is seen as enhancing primary wind-borne dispersal from the parent plant to the ground. However, the consequences of the samara wing of primarily wind-dispersed tree species for post-dispersal processes has not been well studied. We explored whether the presence of this wing in Acer pseudoplatanus either deters or promotes predation after dispersal, either by increasing the time and energy required to predate the seed or by increasing the seed's visibility to predators. We found that wing-removed fruits were preferred, suggesting that the presence of samaras makes seed handling more expensive for granivores. Further, we found that fewer seeds were consumed from treatments that contained the most winged seeds, thus there was no evidence of the samaras making seed finding easier for granivores. We conclude that the presence of the wing may offer an anti-predatory benefit as well as aiding primary dispersal.     

Laccase from Sycamore Maple (Acer pseudoplatanus) Polymerizes Monolignols

Current understanding of the final oxidative steps leading to lignin deposition in trees and other higher plants is limited with respect to what enzymes are involved, where they are localized, how they are transported, and what factors regulate them. With the use of cell suspension cultures of sycamore maple (Acer pseudoplatanus), an in-depth study of laccase, one of the oxidative enzymes possibly responsible for catalyzing the dehydrogenative polymerization of monolignols in the extracellular matrix, was undertaken. The time course for secretion of laccase into suspension culture medium was determined with respect to age and mass of the cells. Laccase was completely separated from peroxidase activity by hydrophobic interaction column chromatography, and its purity was assessed with different types of gel electrophoresis (isoelectric focusing-, native-, and sodium dodecyl sulfate-polyacrylamide gel electrophoresis). Amino acid and glycosyl analyses of the purified enzyme were compared with those reported from previous studies of plant and fungal laccases. The specific activity of laccase toward several common substrates, including monolignols, was determined. Unlike a laccase purified from the Japanese lacquer tree (Rhus vernicifera), laccase from sycamore maple oxidized sinapyl, coniferyl, and p-coumaryl alcohols to form water-insoluble polymers (dehydrogenation polymers).     

Evolutionary correlations of polycyclic shoot growth in Acer (Sapindaceae)

Two strategies have evolved in understory trees in relation to light availability: maximization of light capture and shade tolerance. In the genus Acer, light capture is favored by a suite of traits maximizing twig thickness and leaf size and minimizing the density of branching in the crown. In contrast, shade tolerance is enhanced by minimizing crown area, crown volume, and total leaf area per unit height. Maples with polycyclic shoot growth (i.e., successive flushes of shoot growth separated by a resting phase within the same vegetative period) may benefit from the prolonged growth by growing more and increasing total leaf area; thus we hypothesize that polycyclism is evolutionarily correlated with the suite of traits related to light capture. We tested this hypothesis using different phylogenetic trees to explore correlations between polycyclism and both suites of traits. Polycyclism was correlated with the suite of traits maximizing light capture, suggesting that polycyclic maples are "optimists" (i.e., they make vigorous vertical extensions in rich light) and monocyclic maples are "pessimists" (i.e., they wait in the dark understory until a gap is opened). Both strategies have been described for different floras, and interestingly, polycyclic species recruit over a wider range of environments than the monocyclic species.     

Uridine diphosphate-sugar metabolism by sycamore (Acer pseudoplatanus) cambial tissue

Summary Extracts of sycamore cambial tissue convert UDP-glucose into UDP-glucuronic acid, and the latter into UDP-xylose and UDP-rhamnose. None of the corresponding galactose series of monosaccharides was formed indicating the absence of epimerases, postulated as an important feature of differentiation. UDP-glucose is formed in greater quantities than any of its nucleoside analogues and it is suggested that UDP-glucose plays a more important role in carbohydrate metabolism in this tissue.     

Extracellular laccases and peroxidases from sycamore maple (Acer pseudoplatanus) cell-suspension cultures

We have investigated the abilities of extracellular enzymes from dark-grown cell-suspension cultures of sycamore maple (Acer pseudoplatanus L.) to oxidize monolignols, the precursors for lignin biosynthesis in plants, as well as a variety of other lignin-related compounds. Laccase and peroxidase both exist as a multiplicity of isoenzymes in filtrates of spent culture medium, but their abilities to produce water-insoluble, dehydrogenation polymers (DHPs) from the monolignols (in the presence of hydrogen peroxide for the peroxidase reaction) appear identical whether or not the enzymes are purified from the concentrated filtrates or left in a crude mixture. The patterns of bonds formed in these DHPs are identical to those found in DHPs synthesized using horseradish peroxidase or fungal laccase, and many of these bonds are found in the natural lignins extracted from different plant sources. On the other hand, sycamore maple laccase is very much less active on phenolic substrates containing multiple aromatic rings than is sycamore maple peroxidase. We suggst that whereas laccase may function during the early stages of lignification to polymerize monolignols into oligo-lignols, cell-wall peroxidases may function when H₂O₂ is produced during the later stages of xylem cell development or in response to environmental stresses.Abbreviations DHP dehydrogenation polymer - IEF isoelectric focuring - NMR nuclear magnetic resonance - PAGE polyacrylamide gel electrophoresis The authors wish to thank Dr. Masahiro Samejima (University of Tokyo) for provision of lignin model compounds and Dr. Göran Gellerstadt (Royal Institute of Technology, Sweden) for helpful suggestions regarding stilbene formation and light spectroscopy. Monolignols were prepared by Mr. Nate Weymouth with help from Dr. Herb Morrison (USDA/ARS, Richard B. Russell Research Center, Athens, GA). Thanks also to Ms. Izabella Poppe of the Complex Carbohydrate Research Center (CCRC) for assistance with carbohydrate analyses, and Mr. Vincent Sorrentino for help with the growth of cell-suspension cultures.     

Acclimation to Drought in Acer pseudoplatanus L. (Sycamore) Seedlings

A glasshouse experiment was conducted with well-watered andwater-stressed seedlings of sycamore (Acer pseudoplatanus L.)grown in soil columns. Water was withheld when the seedlingswere 82-d-old. Effects of soil drying on stomatal behaviour,water relations, xylem cavitation, and growth of leaves androots were evaluated. Stomatal conductance declined well before any observable changein bulk leaf water potentials, and was correlated with soilwater status. At seven weeks, osmotic potential had declinedby 0·51 MPa and 0·44 MPa at full and zero turgor,respectively. Drought significantly increased both bulk elasticmodulus and leaf dry weight to turgid weight ratio of water-stressedplants. Drought had no effect on relative water content at zeroturgor. Water cavitation in the xylem was detected as ultrasonic acousticemissions (AE). Water-stressed plants displayed significantlyhigher rates of AE than well-watered plants. Maximum rate ofAE coincided with the minimum level of stomatal conductanceand apparent rehydration of the leaves. Drought caused changes in the root distribution profile andit increased the root weight. The increase in root weight wasmainly due to a substantial shift in assimilates allocated infavour of roots with total biomass being unaffected. Leaf growthwas maintained for six weeks without any significant declinein expansion rate. However, the development of severe waterstress reduced both leaf production and expansion.     

Cellulase and cell differentiation in Acer pseudoplatanus

Summary Homogenates of differentiating xylem and phloem tissue have higher cellulase activities than cambial samples; the highest activity is always found in phloem. Callus tissue, in which no vascular differentiation occurs, contains only low cellulase activity. The results suggest that cellulase is involved in vascular differentiation. Different pH optima of cellulase activity were found: in cambium, xylem and phloem tissue, cellulase activity with an optimum at about pH 5.9 is predominantly membrane-bound; it is sedimentable at 100,000 g and releasable by Triton X-100. The same may be true of activity with an optimum at pH 5.3. Phloem tissue also contains a soluble, cytoplasmic cellulase of high activity at pH 7.1, and xylem tissue contains cytoplasmic cellulase with an optimum at pH 6.5. Low cellulase activity with a pH optimum similar to that of xylem homogenates was found in xylem sap. Cellulase activity in abscission zones increases greatly just before leaf abscission. Abscission zone cellulase has two pH optima, et 5.3 and 5.9; both activities are increased by Triton treatment of homogenates. The possible existence of several different cellulases forming part of a cellulase complex, and the rôle of the enzymes in hydrolysing wall material during cell differentiation are discussed.     

Tapering of xylem conduits and hydraulic limitations in sycamore (Acer pseudoplatanus) trees

Vertical conduit tapering is proposed as an effective mechanism to almost eliminate the increase in hydraulic resistance with increased height. Despite this potential role, very little is known about its changes during ontogeny. Here, conduit tapering and stem morphology of young/small and old/tall individuals of Acer pseudoplatanus in the field, as well as 3-yr-old grafted trees from both age classes, were analysed. The distribution of hydraulic resistance along stems was also determined in a subsample of trees. Substantial conduit tapering was found in small trees (field-grown and grafted from both age classes), whereas values were lower in tall trees, indicating that tapering was a size-related, not an age-related process. Apical conduit diameters were larger in tall trees and were inversely correlated with the degree of tapering. Hydraulic resistance increased less than linearly with distance from the apex. Its scaling against distance was indistinguishable from that predicted from anatomical measurements. Conduit tapering was an effective but partial mechanism of compensation for the increase in hydraulic resistance with tree height. Size-related changes in tapering and in apical conduit diameters may be explained by the combined need to reduce the build-up of hydraulic resistance while minimizing the carbon costs of building vessel walls.     

Seed dormancy in Acer: Endogenous germination inhibitors and dormancy in Acer pseudoplatanus L.

Summary Dormant seeds of Acer pseudoplatanus L. contain two zones of inhibition on paper chromatograms in 10:1:1 as detected by the lettuce and cress seed germination, and the wheat coleoptile bioassays. One zone at R_f 0.6–0.8 was partitioned into ethyl acetate at acid pH and was shown to contain ABA by its behaviour on GLC and isomerization under ultra-violet light. The other zone at R_f 0.9 was detected only in the germination bioassays and was partitioned into ethyl acetate over a range of pH indicating the presence of one or more neutral compounds.The inhibitors present in the embryo of dormant sycamore seeds inhibited the germination of non-dormant sycamore seeds at relatively low concentrations. A comparison with the effects of application of exogenous ABA indicated that endogenous ABA could not solely account for the inhibitory activity of seed extracts, which appeared to be due partly to the presence of ABA and partly to that of neutral compounds present in the embryo. Leaching treatments that removed dormancy led to a decrease in the level of inhibitors present mainly in the basic fraction. The exogenous application of kinetin to dormant sycamore seeds increased germination whereas gibberellic acid had no effect. Similar responses were obtained with lettuce seeds inhibited by the basic fraction of dormant sycamore seeds.It is suggested that an inhibitor-cytokinin interaction may be involved in the dormancy of sycamore seeds.     

Multiple Metal Resistance and Co-resistance in Acer pseudoplatanus L. (Sycamore) Callus Cultures

In vitro growth of Acer pseudoplatanus L. (sycamore) callustissue derived from shoot-tip explants and screened on metal-enrichedmedia was studied in an attempt to identify resistance traitswhich may explain the survival of trees at metal-contaminatedsites. Copper and Cd-resistance traits were identified in celllines originating from trees at a site with a relatively recenthistory of severe contamination by these metals, and Cd- andZn-resistance were identified in cell lines originating frommature trees at a mining spoil site with a much longer historyof exposure to elevated concentrations of these metals. In Zn-resistantcell lines, co-resistance to Ni was also found, even thoughthis metal was not elevated at the study site. This is the firstreport of multiple resistance and co-resistance to metals occurringat the cellular level in trees. The mechanisms of the measuredresistance traits remain unclear, although there was evidenceof reduced Cu and Ni uptake by resistant cell lines. It is concludedthat facultative adaptations allowing acclimation to metal stressmay be particularly significant for survival of mature trees;induction of metal resistance probably occurs in vivo in treesat metal-contaminated sites.Copyright 1995, 1999 Academic Press Acer pseudoplatanus, sycamore, callus, heavy metals, metal resistance, trees     

Separation of membranes from semiprotoplasts of suspension-cultured sycamore maple (Acer pseudoplatanus) cells

Semiprotoplasts were produced from suspension-cultured Acer pseudoplatanus (sycamore maple) cells prior to cell disruption by passing them through a 60 μm nylon screen. Cell membranes from homogenates were separated by ultracentrifugation on linear sucrose density gradients. Samples were collected by gradient fractionation and subcellular fractions were assayed for membrane markers and glycosyl transferase activities. Results of standard marker assays (cytochrome c reductase for endoplas-mic reticulum. uridine and inosine diphosphatases for Golgi. and eosin-5'-maleimide binding for plasma membrane) showed partial separation of these three membrane types. Golgi and plasma membrane markers overlapped in most gradients. Incorporation of ¹⁴C-labeled sugars from UDP-glucose and UDP-xylose into ethanol precipitated polysaccharides was used to detect glucan synthases I & II (glucosyl transferases) and xylosyl transferase activities in Golgi membrane fractions. All three glycosyl transferases overlapped in fractions corresponding to both Golgi and plasma membrane markers, although peak activities for all three occurred in different fractions. More than one peak of glucan synthase I activity was found. Glucan synthase II, associated with ß-l.3 glucan (cullose) synthesis in plasma membranes, was also detected and exhibited a 10-fold stimulation in the presence of Ca²⁺.     

Inflorescence dimorphism,heterodichogamy and thrips pollination in Platycarya strobilacea (Juglandaceae)

Background and Aims

Unlike other taxa in Juglandaceae or in closely related families, which are anemophilous, Platycarya strobilacea has been suggested to be entomophilous. In Juglandaceae, Juglans and Carya show heterodichogamy, a reproductive strategy in which two morphs coexist in a population and undergo synchronous reciprocal sex changes. However, there has been no study focusing on heterodichogamy in the other six or seven genera, including Platycarya.

Methods

Inflorescence architecture, sexual expression and pollination biology were examined in a P. strobilacea population in Japan. Flowering phenology was monitored daily for 24 trees in 2008 and 27 in 2009. Flower visitors and inhabitants were recorded or collected from different sexes and stages.

Key results

The population of P. strobilacea showed heterodichogamous phenology with protogynous and duodichogamous–protandrous morphs. This dimorphism in dichogamy was associated with distinct inflorescence morphologies. Thrips pollination was suggested by the frequent presence of thrips with attached pollen grains, the scarcity of other insect visitors, the synchronicity of thrips number in male spikes with the maturation of female flowers, and morphological characters shared with previously reported thrips-pollinated plants. Male spikes went through two consecutive stages: bright yellow and strong-scented M1 stage, and brownish and little-scented M2 stage. The latter contained more thrips, synchronized better with the receptive stage of female flowers of the reciprocal morph and is probably the main period of pollen export.

Conclusions Platycarya strobilacea

is heterodichogamous and thrips-pollinated, both of which are relatively rare conditions in angiosperms. In male spikes of P. strobilacea, there is probably a temporal decoupling of pollinator attraction and pollen export.     

Xylan synthetase activity in differentiated xylem cells of sycamore trees (Acer pseudoplatanus)

Particulate enzymic preparations obtained from homogenates of differentiated xylem cells isolated from sycamore trees, catalyzed the formation of a radioactive xylan in the presence of UDP-D-[U-¹⁴C]xylose as substrate. The synthesized xylan was not dialyzable through Visking cellophane tubing. Successive extraction with cold water, hot water and 5% NaOH dissolved respectively 15, 5 and 80% of the radioactive polymer. Complete acid hydrolysis of the water-insoluble polysaccharide synthesized from UDP-D-[U-¹⁴C]xylose released all the radioactivity as xylose. -1,4-Xylodextrins, degree of polymerization 2, 3, 4, 5 and 6, were obtained by partial acid hydrolysis (fuming HCl or 0.1 M HCl) of radioactive xylan. The polymer was hydrolysed to xylose, xylobiose and xylotriose by Driselase which contains 1,4- xylanase activities. Methylation and then hydrolysis of the xylan released two methylated sugars which were identified as di-O-methyl[¹⁴C]xylose and tri-O-methyl-[¹⁴C]xylose, suggesting a 14-linked polymer. The linkage was confirmed by periodate oxidation studies. The apparent K_m value of the synthetase for UDP-D-xylose was 0.4 mM. Xylan synthetase activity was not potentiated in the presence of a detergent. The enzymic activity was stimulated by Mg²⁺ and Mn²⁺ ions, although EDTA in the range of concentrations between 0.01 and 1 mM did not affect the reaction rate. It appears that the xylan synthetase system associated with membranes obtained from differentiated xylem cells of sycamore trees may serve for catalyzing the in vivo synthesis of the xylan main chain during the biogenesis of the plant cell wall.     

Characterization of microsatellite markers in sycamore (Acer pseudoplatanus L.)

Sycamore (Acer pseudoplatanus L.) is a tetraploid European hardwood tree species. The reproduction system of the insect‐pollinated trees and patterns of genetic variation are largely unknown. We isolated and characterized eight polymorphic microsatellite markers for Acer pseudoplatanus L. The high degree of polymorphism observed at these markers makes them useful to observe genetic variation patterns at various spatial scales and to analyse gene flow and the mating system. Primers developed for the amplification of microsatellites in A. pseudoplatanus were tested for 21 different species of genus Acer. Amplification products of the expected size were obtained in most cases.     

Competition for nitrogen sources between European beech (Fagus sylvatica) and sycamore maple (Acer pseudoplatanus) seedlings

To investigate the short‐term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter‐specific competition.     

Effect of sucrose starvation on sycamore (Acer pseudoplatanus) cell carbohydrate and Pi status.

The mobilization of stored carbohydrates during sucrose starvation was studied with sycamore (Acer pseudoplatanus) cells. When sucrose was omitted from the nutrient medium, the intracellular sucrose pool decreased rapidly during the first hours of the experiment, whereas the starch content remained practically unchanged. After 10h of sucrose starvation, starch hydrolysis replaced sucrose breakdown. From this moment, the phosphate-ester pool and respiration rate decreased with time. Conversely, the intracellular Pi concentration increased. 31P n.m.r. of intact sycamore cells indicated that, under these conditions, most of the Pi accumulated in the vacuole. These results strongly suggest that starch breakdown, in contrast with sucrose hydrolysis, is not rapid enough to maintain a high cellular metabolism.     

Heterodichogamy and nitidulid beetle pollination in Anaxagorea prinoides, an early divergent Annonaceae

Heterodichogamy in a natural population of an Annonaceae species from the rainforests of French Guiana is described for the first time. Anaxagorea prinoides had bisexual flowers and two floral morphs within the studied population were protogynous. The population under study comprised 7 mature trees belonging to one morph and 12 to the other. Statistical analyses showed that the two morphs were in a 50:50% ratio, and therefore the temporal sexual pattern of heterodichogamy is given. When anthesis of flowers in the male stage ended in one morph, anthesis started with flowers in the female stage in the complementary morph. Approximately 1?h before the end of anthesis in one morph, flowers of the reciprocal morph started to emit a fruit-like scent. The temporal separation of the female and the male stages of the two different morphs lasted only approximately 1?h. Six of the seven identified compounds in the banana-like floral scent were esters and one was an alcohol. The main compounds examined are known to be components of fruit scents. Nitidulidae beetles of the genus Colopterus were the pollinators of A. prinoides and during flowering were maintained within the population of this species. This was not only due to the fact that the beetles remained sheltered in the pollination chamber of the flowers, but also because upon release from individuals of male-stage flowers at the end of flower anthesis they were attracted by the odoriferous female-stage flowers of other individuals of the same population. Heterodichogamy of A. prinoides appears to be a means by which reproductive success is augmented. Attraction of beetle pollinators by ??fruit-imitating?? floral scent is not restricted to species of Anaxagorea, but occurs in many representatives of the Annonaceae.