Effects of microsite light availability on the survival and growth of oak seedlings within a grassland

Effects of microsite light availability on the growth and survival of transplantedQuercus serrata Thunb. seedlings in aMiscanthus sinensis Anderss. grass canopy were investigated by a “plant's eye view” approach. Diffuse site factors, i.e., the fractional transmissions of diffuse photosynthetic photon flux density, were estimated at 15 cm aboveground in 108 microsites where the seedlings grew. Microsite diffuse site factors were significantly different between surviving and dead seedlings during the experiment period from April to October (F_[1,14]=10.9, P<0.01). Relative growth rate of dry weight for individual seedlings positively depended on the diffuse site factors (r²=0.482, P<0.001 in May; r²=0.312, P<0.001 in October). Only 16 seedlings produced their second stem flush within the grass canopy. The ratio of height to dry weight of the second stem flush was significantly higher for the seedlings grew in shady microsites than for those in less shady microsites (r²=0,471, P<0.01 in May). This study suggests that the microsite heterogeneity of light availability is one of the important factors affecting the establishment of tree seedlings in patchy grasslands.     

Seasonal variations of microsite light availability within aMiscanthus sinensis canopy

Photosynthetic photon flux density (PPFD) at 15 cm above the ground was measured at 20 microsites in gaps and grass patches within aMiscanthus sinensis Anderss community at 10 s intervals during 5 days every month from May to September 1989. Microsite light availability, which was characterized by daily total PPFD, sunfleck PPFD (PPFD above a threshold value of 50 or 400 μmol m⁻² s⁻¹) and the diffuse site factor, showed evident seasonal changes, with a marked reduction between June and July due to the rapid growth of the grass canopy. The monthly median value of daily total PPFD among the microsites decreased from 10.3 mol m⁻² day⁻¹ in May to 0.77 mol m⁻² day⁻¹ in September, with a reduction in the diffuse site factor from 31 to 4%. During the summer, the median value of the total time of sunflecks exceeding 50 μmol m⁻² s⁻¹ contributed 7–18% of measurement time, but the contribution of these sunflecks to daily total PPFD ranged from 29 to 59%. There was considerable microsite variation in light availability throughout the measurement period. Rank correlation analysis revealed that some microsites, such as those in gaps, consistently received more total PPFD, more sunfleck PPFD and had a higher diffuse site factor than those in grass patches. The diffuse site factor had a linearly positive relationship with daily total PPFD and total sunfleck PPFD for the 20 microsites during the measurement period, confirming that the diffuse site factor is a useful index for microsite light availability withinM. sinensis canopies.     

Comparative structural study ofQuercus serrata andQ. acutissima formed byPisolithus tinctorius andHebeloma cylindrosporum

Ectomycorrhizas were synthesized in pots and growth pouches betweenQuercus serrata, Q. acutissima, and two ectomycorrhizal fungi,Pisolithus tinctorius andHebeloma cylindrosporum. Root morphology and the structure of the mantle and Hartig net were compared using light, fluorescence, scanning and transmission electron microscopy.P. tinctorius initially colonized root cap cells, and eventually produced a highly branched lateral root system with a complete mantle, whereasH. cylindrosporum promoted root elongation with few hyphae on the root apex surface indicating that interaction between roots differs with fungal species. Hartig net structure and hyphal inclusions varied between all the combinations tested. There were structural differences between mycorrhizas ofH. cylindrosporum/Q. acutissima grown in soil and growth pouches, which indicate that the growth pouch environment can induce artefacts in roots. Fruit bodies ofH. cylindrosporum developed in pots withQ. acutissima. AlthoughP. tinctorius has been used to inoculate oak seedlings in the nursery, results of this study indicate thatH. cylindrosporum may also be an effective ectomycorrhizal fungus forQ. serrata andQ. acutissima.     

Microsite variation in light availability and seedling growth ofQuercus serrata in a temperate pine forest

Spatial heterogeneity in light availability for tree seedlings under the canopy of a temperate pine forest was studied. Six-day measurements at 10-s intervals revealed a great variety in the temporal patterns of photosynthetic photon flux density (PPFD) and histograms among observation days and microsites; mean daily total PPFD relative to full sun varied from 1.5% to 10.4% depending on the microsites. The occurrence and duration of PPFD above 80 μmol m⁻² s⁻¹, which might reflect sunfleck activity, varied greatly among the microsites. However, several simple empirical relationships were found between some parameters characterizing microsite light availability and sunfleck activity; the diffuse site factor was correlated well with other parameters, including daily total PPFD, daily totals and daily summed durations of high PPFD above any examined threshold level, and its contribution to daily total PPFD. Diffuse site factors which were obtained for 700 microsites within an area of 28 m² on three different occasions during the growing season showed high correlations within the microsite. Based on the regressed relationship between the relative growth rate of current-year seedlings ofQuercus serrata and the microsite diffuse site factor and the results of area-survey measurement of the diffuse site factor, an estimation was made of the abundance of potential ‘safe-sites’ for seedling growth of the species; the ‘safe sites’ were estimated to cover 40% and 0% of the total area of the sunny and shady sites of the forest, respectively.     

Fluctuation of photosynthetic photon flux density within aMiscanthus sinensis canopy

Detailed measurements of diurnal variations in photosynthetic photon flux density (PPFD) were made at seven locations within the canopy of aMiscanthus sinensis grassland to evaluate the light conditions of microsites for heliophilic tree seedlings. Multiple regression analysis revealed that the short-term light fluctuation on a clear day was highly dependent on the wind speed and solar elevation angle, whereas on a cloudy day it was mainly determined by the PPFD incident from above the canopy. The relative PPFD at 40 cm aboveground varied from 0.065 to 0.252, depending on sky conditions and the sensor's position in relation to clumped patches ofM. sinensis. On a clear day, the proportion of PPFD readings above 100 μmol·m⁻²·s⁻¹ contributed by sunflecks ranged between 25.4% and 82.0%. Computer simulation showed that the contribution of sunflecks to the daily carbon gain ofQuercus serrata seedlings may range from 11% to 65%. The mean relative PPFD measured under diffuse light conditions was linearly related to the daily total PPFD and the daily carbon gain by single leaves ofQ. serrata seedlings. This suggests that the relative PPFD under diffuse light conditions provides an appropriate measure of site-specific light availability within a grass canopy.     

Callus proliferation from the protoplasts of embryogenic cells of Quercus serrata

Embryogenic culture was induced from the immature embryos of Quercus serrata using Marashige and Skoog's medium (MS) containing 0.1 M each of 2,4-d and BAP, and subcultured for seven months before isolation of protoplasts by using 1% Cellulase RS in 0.6 M mannitol solution. Efficient colony formation was obtained when protoplasts were cultured in a liquid MS medium containing 0.6 M mannitol, 3% sucrose and combination of 0.1 M or 1 M each of 2,4-d and BAP. Excluding ammonium nitrate from the MS medium resulted in the decrease of the percentage of colony formation. From colonies, both agar culture and liquid culture were sustained in the MS media without mannitol containing no plant growth regulator, or containing 0.1 M of BAP in combination with 0.1 M or 1 M of 2,4-d.Abbreviations BAP 6-benzylaminopurine - 2,4-d 2,4-dichlorophenoxyacetic acid - MS medium after Murashige & Skoog (1962).     

Species diversity and primary productivity inMiscanthus sinensis grasslands

In an old field grassland dominated byMiscanthus sinensis Anderss. the community structures, phytomass, dominance ofM. sinensis and species diversity were measured. Species and life form composition of the stand were characterized by higher percentages of therophytes, woody and shrubby species, liana and alien species. From May so September in 1982, all the aboveground parts were harvested from each of the four quadrats (2 m×2 m) once a month. Seasonal peak of aboveground phytomass, in September, was 1027 g d.w.m⁻² to whichM. sinensis contributed as much as 96.5%. With the progress of the growing season,M. sinensis became increasingly important both in stand phytomass and in dominance, whereas species diversity based on the dry weight contributions of constituent species decreased. Our analysis of these seasonal trends showed that the diversity was largely a function of dominance of the most important species, rather than that of stand phytomass or productivity. The simultaneous measurements of 20 quadrats in late August 1983, also supported the above conclusion.     

Production-dependent reproductive allocation of a tall tree speciesQuercus serrata

The reproductive allocation (allocation of net production to acorns) of a tall tree speciesQuercus serrata was estimated by a method combining branch diameter distribution and the sampling of acorns per branch. Acorn production per branch of 0.5-cm diameter significantly varied among individuals and years, but not significantly across tree size. Leaf production per branch of 0.5-cm diameter had little annual fluctuation, while that of acorns fluctuated about 6 times between the maximum and minimum. The reproductive allocation was regressed against vegetative allocation, following the general model of klinkhameret al. (1992); the reproduction had a beginning threshold and reproductive allocation increased drastically at relatively young stage after the tree began reproduction. With its small threshold production, coppicing traits and large maximum size,Q. serrata seemed adaptive to infrequent but large-scale disturbances like fires.     

Phytoclimatic characterization ofQuercus frainetto Ten. stands in peninsular Italy

The floristic differentiation of the deciduousQuercus frainetto forests along the climatic gradients of Apennine Italy has been studied. The ecological amplitude of this oak, and the bioclimatological relationships here assessed, suggest potentiality for the growth ofQ. frainetto-rich communities as broad zonal vegetation belt, ranging from NW to SE along the W side of peninsular Italy. Strong floristical and ecological similarities to the balcanic stands are described.The status of real vegetation belt for the ItalianQuercion frainetto s.l. is here emphasized.     

Growth and shoot: root ratio of seedlings in relation to nutrient availability

The influence of mineral nutrient availability, light intensity and CO₂ on growth and shoot:root ratio in young plants is reviewed. Special emphasis in this evaluation is given to data from laboratory experiments with small Betula pendula plants, in which the concept of steady-state nutrition has been applied.Three distinctly different dry matter allocation patterns were observed when growth was limited by the availability of mineral nutrients: 1, Root growth was favoured when N, P or S were the major growth constraints. 2, The opposite pattern obtained when K, Mg and Mn restricted growth. 3, Shortage of Ca, Fe and Zn had almost no effect on the shoot:root ratio. The light regime had no effect on dry matter allocation except at very low photon flux densities (< 6.5 mol m^-2 day^-1), in which a small decrease in the root fraction was observed. Shortage of CO₂, on the other hand, strongly decreased root development, while an increase of the atmospheric CO₂ concentration had no influence on dry matter partitioning. An increased allocation of dry matter to below-ground parts was associated with an increased amount of starch in the tissues. Depletion of the carbohydrate stores occurred under all conditions in which root development was inhibited. It is concluded that the internal balance between labile nitrogen and carbon in the root and the shoot system determines how dry matter is being partitioned in the plant. The consistency of this statement with literature data and existing models for shoot:root regulation is examined.     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

Coleoptiles are gravi-guiding systems vital for gravi-insensitive shoots of germinating grass seedlings

Surgical removal of the coleoptiles from germinating seedlings of Secale cereale L., Avena sativa L., and Zea mays L. leads to a total loss of gravitropic response of the shoot without affecting growth. The principal function of coleoptiles is therefore to guide the gravi-insensitive shoot to the soil surface.Dedicated to Prof. Andreas Sievers on the occasion of his retirement     

Spatial heterogeneity of photosynthetic photon flux density in the canopy ofMiscanthus sinensis

Spatial variation in photosynthetic photon flux density (PPFD) was investigated in detail at different heights within the canopy of aMiscanthus sinensis grassland to evaluate the light environment of microsites for establishment of heliophilic tree seedlings. Highly heterogeneous patterns of light distribution were revealed within the apparently uniform grass canopies, especially under direct light. The frequency distribution patterns of relative PPFD (RPFD) were compared among different solar and sky conditions. With increasing height in the canopy, the mean RPFD value and standard deviation (SD) increased, while the skewness and kurtosis of the distribution decreased. The mean RPFD and SD were higher, especially at higher solar elevation angles, under direct light than those under diffuse light conditions. The frequency distribution of RPFD was more platykurtic under direct light and at higher solar elevation angles.     

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size

Summary We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.     

Population dynamics and growth patterns for a cohort of northern red oak (Quercus rubra) seedlings

Summary I studied the survival and development of a 1986 cohort of northern red oak (Quercus rubra L.) seedlings growing under a variety of overstory and microsite conditions in a northern hardwood forest dominated by northern red oak, red maple (Acer rubrum L.) paper birch (Betula papyrifera Marsh.), and scattered white pine (Pinus strobus L.). Fifty naturally regenerating seedlings of oak were randomly selected in each of three canopy classes: no overstory, partial overstory, and complete overstory. Growth and mortality were measured for six years. Seedling height growth decreased with overstory density, with less growth evident with even a partial overstory. Seedling survival also declined with overstory density and depended on microtopography to a lesser extent. After six years, 92% of the seedlings survived in the open, compared to 54% under the partial overstory, and 36% under the complete overstory. The open environment, in which woody and herbaceous regrowth formed a low canopy reducing light intensities to about 50% of full sunlight, provided a favorable site for the growth and survival of northern red oak.     

Guild structures of seed insects in relation to acorn development in two oak species

We investigated patterns of acorn growth in Quercus variabilis Blume and Quercus serrata Thunb., seasonal trends in emergence of seed insects found in the acorns, oviposition periods of the insects and falling periods of insect-infested acorns. In Q. variabilis, two insect guilds were associated with acorn development: (i) the immature acorn feeding (IAF) guild [Curculionidae sp., Poecilips cardamomi (Schaufuss), and Characoma ruficirra (Hampson)]; and (ii) the mature acorn feeding (MAF) guild [Curculio robustus (Roelofs), Curculio sikkimensis (Heller), and Cydia glandicolana (Danilevsky)]. In Q. serrata, there were three guilds: (i) the pistillate flower feeding (PFF) guild (cynipid wasp); (ii) the IAF guild [sap absorption by Mechoris ursulus (Roelofs)]; and (iii) the MAF guild (M. ursulus, C. sikkimensis, Cydia danilevskyi (Kuznetzov), C. glandicolana and Autostichidae sp.). The succession of guilds during acorn development may be a consequence of the use by different species of the limited food resource. The lack of a PFF guild in Q. variabilis that was found in our field site would have a positive effect on IAF guilds in utilizing the acorns.     

Acclimation to light in seedlings of <Emphasis Type="Italic">Quercus petraea</Emphasis> (Mattuschka) Liebl. and <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd. planted along a forest-edge gradient

Photosynthetic acclimation of two co-occurring deciduous oaks (Quercus petraea and Quercus pyrenaica) to a natural light gradient was studied during one growing season. In the spring of 2003, 90 seedlings per species were planted along a transect resulting from a dense Pinus sylvestris stand, an adjacent thinned area and a 10-m-wide firebreak (16.5–60.9% Global Site Factor (GSF)). In two dates of the following summer, we measured leaf gas exchange, carboxylation efficiency (CE), chlorophyll and nitrogen content, light–response curves of chlorophyll a fluorescence parameters, and leaf mass per area (LMA). Summer was mild, as evidenced by leaf predawn water potential (Ψ_pd), which reduced the interactive effect of water stress on the response of seedlings to light. Q. pyrenaica had higher LMA, CE, stomatal conductance (g _{s max}) and photosynthesis per unit area than Q. petraea at all growth irradiances. , LMA, g _{s max} and electron transport rate (ETR) all increased with light availability (GSF) in a similar fashion in both species. Light had also a clear effect on the organization of Photosystem II (PS II), as deduced by chlorophyll a fluorescence curves. Chlorophyll concentration (Chl_m) decreased with increasing light availability in Q. pyrenaica but it did not in Q. petraea. Seedlings of Q. petraea acclimated to higher irradiances showed a greater non-photochemical quenching (NPQ) than those of Q. pyrenaica. This suggests a higher susceptibility to high light in Q. petraea, which would be consistent with a better adaptation to shade, inferred from the lower LMA or the lower rate of photosynthesis.     

Pathogenicity of fungi isolated from the larvae of the mangrove crab,Scylla serrata, in Indonesia

Heavy mortality reaching almost 100% occurred in the larvae of the mangrove crab,Scylla serrata, from July to December 1997 at the hatchery of Gondol Research Station for Coastal Fisheries in Bali, Indonesia. Mortality was observed in larvae after hatching from the eggs. The affected larvae were whitish and filled with numerous aseptate hyphae. Three fungi belonging to the order Lagenidiales,Lagenidium callinectes, Haliphthoros milfordensis, andHalocrusticida sp., were isolated from the infected larvae. Pathogenicity tests of the infected fungi against the larvae of mangrove crab demonstrated that all isolates were pathogenic.     

Growth responses of rice seedlings to triacontanol in light and dark

Triacontanol, a 30-carbon primary alcohol, applied in nutrient culture solutions to rice (Oryza sativa L.) seedlings at 2.3×10^-8 M (10 g/l), caused an increase in dry weight and leaf area of the whole plants. The response could be observed as early as 3 h of treatment. It was observed at relatively high and low light intensities as well as in the dark where control plants lost but triacontanol-treated plants gained in dry weight. The dry weight gain in the dark was, however, eliminated by removing CO₂ from the atmosphere. Triacontanol-treated plants also increased their content of Kjeldahl-N and contained 30% more total N per plant than controls after 6 h in the dark.     

Influences of nitrogen load on the growth and photosynthetic responses of <Emphasis Type="Italic">Quercus serrata</Emphasis> seedlings to O<Subscript>3</Subscript>

The objectives of this study were to clarify the influences of nitrogen (N) load on the growth and photosynthetic responses of Quercus serrata seedlings to O₃ and to obtain basic data for evaluating the critical levels of O₃ for protecting Q. serrata forests in Japan. The effects of O₃ and/or N load on growth and photosynthetic activity of Q. serrata seedlings were investigated during the two growing seasons. Two-year-old seedlings were assigned to 12 experimental treatments, which were comprised of the combination of four gas treatments (charcoal-filtered air and three levels of O₃ at 1.0, 1.5 and 2.0 times ambient concentration) and three N treatments (0, 20 and 50 kg ha⁻¹ year⁻¹). During the second growing season, no significant interactive effects of O₃ and N load on the growth and net photosynthetic rate of the seedlings were detected. Threrfore, we concluded that N supply to the soil at ≤50 kg ha⁻¹ year⁻¹ does not significantly influence the growth and photosynthetic responses of Q. serrata seedlings to O₃. Based on the O₃ exposure-response relationships for the whole-plant growth of the seedlings, the critical level of O₃ for Q. serrata was estimated to be approximately 36 nmol mol⁻¹ as the average 15-h O₃ concentration during the one growing season.