Small-scale patterns of species richness in Swedish semi-natural grasslands: the effects of community species pools

Temperate semi-natural grasslands are known for their high plant species richness at small spatial scales. We examined the variation in small-scale species richness in a sample of 63 sites from Swedish semi-natural grasslands, located as fragments in the modem landscape dominated by forest and agricultural land. Data were obtained from two spatial scales at each site. 1 dm² and 4 m². Using an analysis based on a Monte Carlo simulation, we found support for the species-pool hypothesis: a high species richness at the I dm- scale was associated with high species richness at the 4 m2 scale. The conclusion from this pattern analysis would, however, be considerably strengthened if we could reduce the likelihood that other mechanisms than sampling from species pools of unequal size influence the pattern of small-scale species richness. Additional analyses were made in order to identify such mechanisms. We examined whether four putative key traits: seed size, seed production, plant size and reproductive allocation were different among species at comparatively species-rich vs species-poor I dm' plots. We found only a little evidence for such differences. There was a weak tendency that species in the plots with high species richness possessed larger (and fewer) seeds than species from species-poor plots. Our results are congruent with the main prediction of the species pool model: variation in small-scale species richness (1 dm²- scale) is basically a result of sampling from unequally sized community species pools (4 m² scale). Variation in species richness between the 4 nr semi-natural grassland "patches" may thus be sought for among mechanisms operating al larger spatial scales than 4 m². We briefly discuss such mechanisms, based on other studies performed in the same study area.     

Short-term effects of simulated environmental change on phenology, leaf traits, and shoot growth of alpine plants on a temperate mountain, northern Japan

In order to assess the responses of circumpolar and semicircumpolar plants growing around their southern distribution margins to artificial warming, we set up 11 open-top chambers (OTCs) on a fell-field (1680 m a.s.l.) in the Taisetsu Mountains, northern Japan. The OTCs increased mean air temperature by 1.3°C through the growing season (June–September) and extended the length of the growing season. We examined phenology and leaf traits of plants in the OTCs and control plots during the first season under artificial warming treatment using two deciduous and three evergreen species. Ledum palustre (evergreen shrub), Vaccinium uliginosum , and Arctous alpinus (deciduous shrubs) showed earlier leaf emergence and/or flowering in the OTCs. Deciduous shrubs had longer individual leaf longevity and an extended foliage period in the OTCs than in the control plots. There were no significant differences in specific leaf area and leaf size for many species between the OTCs and the control plots. Vaccinium vitis-idaea (evergreen shrub), L. palustre, A. alpinus , and Empetrum nigrum (evergreen shrub) had lower leaf nitrogen concentration in the OTCs than in the control plots, whereas it was higher in V. uliginosum . Only E. nigrum showed larger annual shoot growth in the OTCs. No clear differences in response to the warming effect were detected between evergreen and deciduous species in the first season. Circumpolar plants growing in temperate alpine regions may be more affected by season length rather than temperature itself.     

Floristic composition, structure, and dynamics of an upper montane rain forest in Southern Ecuador

Two 1 ha plots of undisturbed upper montane rain forest in southern Ecuador were sampled for all trees with a dbh ≥ 5 cm. An extraordinarily high α-diversity for a forest near treeline is described. The "non-ridge forest" plot at 2900 m elevation has 75 species and 28 families ha^-1 and is believed to represent an advanced stage of succession whereas, the "ridge forest" plot at 2700 m elevation with 90 species and 38 families ha^-1 has a more rapid turn-over rate probably due to a more unstable environment. Downslope forces (soil creep) are discussed as a possible cause for 32–44% of all trees being inclined more than 30° in the moderately steep terrain (average slope 20° in both study plots). The plot of "non-ridge forest" is characterized by a much greater biomass whether expressed as basal area (44 m² versus 15 m²) or stem volume (214 m³ versus 52 m³), while the density is equally high (2310 versus 2090 trees ha^-1) in both plots. Families with Family Importance Value >25 / ha^-1 are Clusiaceae, Cunoniaceae, Melastomataceae, Myrsinaceae, and Ternstroemiaceae.     

Responses of alpine shrubs to simulated environmental change during three years in the mid-latitude mountain, northern Japan

Effects of artificial warming on phenology, individual leaf traits, vegetative growth, and reproduction of five alpine species (two deciduous and three evergreen shrubs) were investigated during three years in the mid-latitude alpine, northern Japan. Eleven open-top chambers (OTCs) were set up on a fellfield (1680 m a. s. l.) in the Taisetsu Mountains by which air temperature at plant height was increased by ca 2°C. Vaccinium uliginosum (deciduous shrub) showed earlier leaf emergence in every season and earlier flowering only in the first season in the OTCs. By contrast, acceleration of leaf emergence in the OTCs was not clear for other species, i.e. Arctous alpinus (deciduous shrub). Ledum palustre. V. vitis-idaea , and Empetrum nigrum (evergreen shrub). Both deciduous species showed longer leaf life-span in the OTCs every season. All evergreen species had higher leaf survival rates in the OTCs. indicating extension of leaf life-span. Leaf nitrogen concentration and leaf mass per unit leaf area (mg cm ⁻²) generally tended to decrease in the OTCs. Relationships between the individual leaf traits and cumulative air temperature during the leaf developing period were not clear. Total leaf production during the three seasons increased in the OTCs in A. alpinus. L. palustre. V. vitis-idaea , and E. nigrum. All evergreen shrubs showed larger shoot growth in the OTCs but both deciduous shrubs did not show significant changes. In contrast to the vegetative growth, deciduous shrubs produced more flowers in the OTCs. Fruit production was not influenced by the OTCs for all species. The extension of photosynthetic period in the OTCs may contribute to the larger vegetative growth or flower production.     

Effect of fire on the topsoil seed banks of rehabilitated bauxite mine sites in the jarrah forest of Western Australia

Summary Germinable seed stores of 5- and 8-year-old rehabilitated bauxite mine pits in south-west Western Australia were assessed before and after burning. These seed stores were compared to those of adjacent unmined Jarrah ( Eucalyptus marginata ) forest, to identify at what age fire can be reintroduced, in order to measure restoration success and reduce fire hazard. Soils were sampled in early summer (before fire) and late autumn (after fire). Before fire, the mean topsoil seed bank of 5-year-old sites was 2121 seeds per m² while 8-year-old sites had a mean of 1520 seeds per m². Only the 5-year-old sites were significantly different from the forest mean of 1478 seeds per m² for the same season. After summer burns (and possibly due to seasonal effects) topsoil seed banks of rehabilitated areas (sampled in autumn) decreased by an average of 53 per cent. Topsoil seed banks of 5–8-year-old sites were resistant to lower intensity burns, with 362 seeds per m² of native species surviving mild burns and 108 seeds per m² of native species surviving after an intense summer fire. The topsoil seed reserve of 5–8-year-old rehabilitated areas had a high proportion of annual weed species while the forest sites had high levels of subshrubs and native annuals. Low-intensity burns did not alter the composition of life-forms in the soil seed bank, while intense burns favoured annual weed and shrub species. The results indicate that it is not appropriate to introduce fire to rehabilitated areas before 8 years, due to limited fuel reduction benefits and possible adverse effects on obligate seeding species. The large proportion of weed species in the soil seed bank of young rehabilitated areas is a concern, and remains a major consideration for future disturbance of these areas.     

Fruit and seed ecology of wild robusta coffee (Coffea canephora Froehner) in Kibale National Park, Uganda

Coffee fruit production, frugivorous activities and seed longevity were investigated during two seasons. The average densities of fruiting coffee trees during the first sampling period ranged from four to seven per 25 m² and during the second sampling period, four fruiting trees per 25 m² were estimated. Fruiting coffee trees were significantly less than the non-fruiting trees ( P < 0.05, t -test). Periods of maximum coffee fruit fall coincided with those of lowest rainfall. Coffee fruit yield per tree from ground collections was 7–40 m⁻² in 1992 and 16–38 m⁻² in 1993/1994. Split coffee berries ranged between 71.7% and 83.6% of all fruits collected from the forest floor in the 1993/94 sampling period. Coffee seeds collected from the forest floor were mostly undamaged. Black and white colobus and redtail monkeys were observed to feed on ripe coffee fruits but did not crush the seeds. Coffee seed viability declined rapidly during storage. Seeds left on the forest floor survived longest, those stored under laboratory conditions lost viability fastest, and those in cold storage showed intermediate longevity.     

Two ways of trapping seeds in alpine environments, Lapland, Sweden

Seed dispersal in two habitats was investigated in the mid-alpine Latnjajaure valley, in northernmost Swedish Lapland. The seed rain size was measured using artificial seed traps and natural snowbed at a heath and a meadow site, c. 1000 m a.s.l. The average seed rain size trapped in the snowbeds varied from 18 seeds m^-2 at the heath site, to 96 seeds m-² at the meadow site. On average, the heath trap station had 177 seeds m-² and the meadow station, 218 seeds m². At each site, the vegetation was inventoried within a circular area with a radius of 20 m. Overall, the species assembly in the seed rain reflected the surrounding vegetation. However, there were additional species in the seed rain as well. At the seed trap station on the wind-exposed heath, 78% of the seeds originated from sites outside the homogeneous vegetation around the trap. These seeds were dispersed from locations at least 150 m from the trap site. By trapping a higher number of species than artificial traps and a higher number of seeds per unit area, snowbeds served as effective 'seed traps'. However, the location of the sampling spot in the snowbed is crucial due to a decrease in seed density towards the edge. In this study, with the aim of trapping the main dispersing species in the area, the optimal artificial seed trap size appears to be - 1.5–2.0 m². There was also a positive correlation between the mean July and August temperature, the seed rain size, and the number of species found. Thus, through brief visits to remote locations, the study of seed dispersal can be accomplished effectively when snowbeds are present on the landscape.     

Plant species diversity in an old, traditionally managed hay meadow compared to abandoned hay meadows in southwest Norway

A chronosequence, representing a successional series, was used for the comparison of a hay meadow site managed in an old traditional way for at least a hundred years, and hay meadow sites abandoned for about 10, 20 and 30 years, respectively. Old traditional management included grazing early and late in the growing season, mowing in August and light or no fertilizing. The tree cover was the most important factor deciding the composition of vegetation. Time since abandonment was not completely correlated to tree cover, as some plots had a dense canopy and others were situated in the open. The total species number decreased with number of years since abandonment in plots > 0.001 m²and <100 m². The highest species number in 1 m² plots was recorded in the managed site, with 38 species of phanerogams. Fourtyeight % of the indicators of traditional management present in the managed site was recorded in the site which had been abandoned for 30 years. Frequency — log area curves made it possible to group species according to persistence in the sward. As a result, a group of functional indicators of rare hay meadows in the region was distinguished.     

Effects of experimental small-scale grassland fragmentation on spatial distribution, density, and persistence of ant nests

Abstract.  1. Grassland fragmentation is expected to influence the abundance of different invertebrate species to a different extent. Fragmentation-related effects are of particular importance in species that interact with many other species.
2. The density and spatial distribution of nests of 15 ant species in experimentally fragmented calcareous grasslands at three sites in the Northern Swiss Jura mountains were examined. Fragments of different size (0.25 m², 2.25 m², and 20.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland.
3. Three years after initiation of the experiment, ant nest density did not differ between fragments and control plots. Six years after initiation of the experiment, however, ant nest density and forager abundance were higher in large fragments than in large control plots. Ant nests tended to occur more frequently along the edge of fragments than in the core area. Persistence time of nests of the most abundant species, Lasius paralienus , tended to be shorter in fragments than in control plots. Furthermore, persistence time was longer in nests situated close to the fragment edge than in nests in the core area.
4. Effects on nest density, edge effects on the spatial distribution of nests, and the relationships between nest density and environmental factors were more pronounced when only nests of L. paralienus were considered. The implications of these findings for plant and other invertebrate species are discussed.     

Relationship between seed and clonal growth in the reproduction of Carex rugulosa Kük. in riverside meadows

Carex rugulosa Kük. develops riverside vegetation in brackish marshes along the lower parts of rivers. Because the rhizomatous ramets grow densely (590–950/m²) in its developed population, it is expected that seedlings cannot get enough light to grow if seeds germinate in crowded ramets. We studied the reproductive contribution of seed in a marshy meadow of the Ohashi River, Japan. The number of seeds produced was 6900–14 200/m² in 2002. The number of buried seeds in the following germination season differed among sites from 1190 to 2690/m². No seedlings were observed at plots where the ramets from rhizomes grew densely. In contrast, seedlings appeared in plots where all rhizomatous ramets were artificially cut. The number of seedlings corresponded to 17.5–39.5% of that of buried seeds. All these seedlings were submerged and died when it rained heavily. In the genotype analysis using allozyme detection, one of the phosphoglucoisomerase loci ( Pgi-2 ) indicated a pair of alleles. Among eight plots (each plot was 50 or 60-cm square), four showed an excess of heterozygotes and two showed homozygote domination in the Pgi-2 genotype. These results suggest that: (i) seed germination is suppressed in crowded ramets; (ii) seedling survival is severely reduced by inundation; and (iii) the population is usually maintained by clonal growth. As germination is induced in open areas, it is hypothesized that the main role of seed is recovery after vegetation decline caused by environmental stresses or colonization to other areas without dense vegetation.     

Tree population dynamics in Kibale National Park, Uganda 1975–1998

Changes in species composition, stem abundance, and basal area of trees taller than or equal to 10 m in a medium altitude tropical rain forest at the Ngogo study area, Kibale National Park, Uganda are described for the period between 1975 and 1998 ( n  = 23 years). Two enumeration episodes were conducted in 263 plots of 5 m by 50 m during 1975–80 and 1997–98. During this period, species richness decreased by 3% (from 92 to 89). Species diversity (H') also declined slightly from 2.97 to 2.86. The number of trees recorded in the sample plots decreased by 8% (from 2545 to 2329), while basal area decreased from 49.48 m² ha⁻¹ to 48.68 m² ha^−1. However, stem abundance and basal area increased for some species.     

Patterns of carbon allocation above ground in a deciduous (Vaccinium uliginosum) and an evergreen (Vaccinium vitis-idaea) dwarf shrub

The carbon allocation to current-year shoots of the deciduous Vaccinium uliginosum L. and the evergreen V. vitis-idaea L. was studied in a field experiment using ¹⁴C. During the first week after labelling, 0–50% and 30–80% of the initially assimilated ¹⁴C was lost in V. vitis-idaea and V. uliginosum respectively. Later on, the losses were smaller. After leaf fall in V. uliginosum , 30, 10 and 8% of the initially assimilated ¹⁴C was recovered in the abscised leaves, in plants labelled 1 July, 1 August and 1 September, respectively. The amounts found in the old V. vitis-idaea leaves the year after labelling were 33, 20 and 10%. Only traces of past-year assimilates were found in the current-year V. vitis-idaea leaves, while it was estimated that the V. uliginosum leaves contained 10–15% of the past-year label. It is concluded that V. vitis-idaea is mainly dependent on early summer assimilates - produced by leaves that have overwintered – for the current year shoot growth, while past-years' assimilates probably make an important contribution to the leaf expansion in V. uliginosum. When fruits occurred, a large fraction of the ¹⁴C assimilates was allocated to them.     

Persistence of vascular plants in a Norwegian boreal coniferous forest

Persistence, the tendency of a species to remain in its original position and not to colonize new sites, is studied for the most abundant forest vascular plants (25 species in spruce forest and 7 in pine forest) in Solhomfjell, Gjerstad, S Norway Data sets included presence/absence in 199 meso plots (1 m²) and 3184 subplots (1/16 m²), analyzed over a 5 yr interval, and a subset of 50 meso plots and 800 subplots, analyzed for six consecutive years Relationships between species variables (seedling frequency and mobility rate compiled from the literature, and cover and abundance means in the study area) are studied, and related to species optima along ecologically interpreted DCA ordination axes Vascular plant mobility may increase towards nutrient-poor sites Dominance in the boreal forest floor is mostly by clonal species Persistence was calculated for different temporal (1-5 yr) and spatial (1/16 and I m²) scales Persistence patterns in the spruce and pine forests were similar, but persistence decreased towards the xeric pine forests One main component of variation in persistence was demonstrated by PCA analyses the absolute level of persistence, which is related to seedling recruitment vs clonal growth, and within clonal species to ramet longevity, abundance, mobility, growth pattern and mode of surviving the infavourable season Minor components of variation in persistence were related to spatial scale and temporal scale Persistence characteristics were species-specific and varied little between years Numerous species characteristics were relevant to interpretation of variation in persistence, indicating a continuous, multidimensional variation in life history characteristics     

Dissecting components of population-level variation in seed production and the evolution of masting behavior

Theory predicts that at higher trophic levels species are especially vulnerable to habitat fragmentation due to small population size and dependence on particular prey species. Using aphids as model organism, we tested the hypothesis that herbivore abundance increases in fragmented habitats as a result of reduced predator and parasitoid densities. In a 3 year-study, we examined the population dynamics of aphids with respect to host plant abundance and ant nest density in experimentally fragmented calcareous grasslands at two sites in the northern Jura mountains. Fragments of different size (area: 20.25 m², 2.25 m² and 0.25 m²) were isolated by a 5 m wide strip of frequently mown vegetation and corresponding control plots were situated in the adjacent undisturbed grassland. Aphid density was higher in fragments than in control plots. This was a combined result of a higher frequency of aphid-infested plants and larger aphid colonies in fragments than in control plots. Furthermore, a larger proportion of aphid colonies was ant-attended in fragments than in control plots. Aphid colonies were also more frequently visited by ants in fragments than in control plots in one of the 3 years. Parasitoid pressure on aphids was not influenced by the experimental fragmentation. Neither were aphid species richness and diversity affected by the fragmentation. Our study shows that even small-scale habitat fragmentation can have profound effects on the abundance of herbivorous insects. The effect on aphid density was consistent over 3 years and two sites with slightly different aphid communities.     

The forest of Vorsø, Denmark: Succession towards a natural, decidous boreal forest influenced by breeding cormorants

The forest on the strictly protected island Vorsø in Horsens Fjord has since 1952 been examined every ten years to discover the number, basal area, wood volume and aboveground biomass of the species. This paper describes the results from 1982 and the development up to that. The dominating species are Fraxinus excelsior, Ulmus glabra and Fagus sylvatica. The two forests, "Vesterskov" (6.09 ha) and "Østerskov" (2.48 ha) respectively, were, in 1982, still immature. Their number of stems per ha were: 756 and 509; basal area: 38 m² and 45 m². volume: 544 m³ and 615 m³; biomass 207 t and 268 t. The annual increase 1972–1982 was respectively 8.8 m³/ha and 0.3 m³/ha. The impact of the cormorant ( Phalacrocorax carbo sinensis ) is especially in "Østerskov" increasing, since the colony here was established in 1976. The succession of the forest is discussed in relation to the disturbance by the cormorants.     

Structure, floristic composition, and vegetation forming factors of three vegetation types in Senegal

Six 1 ha plots were established in a coastal savanna, called Fathala Forest, in Delta du Saloum National Park, Senegal. Two plots were placed in woodland, two in wooded grassland, and two in transition woodland in order to describe structure and floristic composition of the vegetation. All trees ≥ 5 cm dbh were sampled. The three selected vegetation types showed distinct differences in structure as well as in species composition. Woodland had high density (440–449 individuals per ha), many small trees, and high basal area (13.4 m²per ha). Transition woodland was characterised by low density (54–118 individuals per ha) but many large trees and a relatively large basal area (8.6–12.8 m² per ha). Wooded grassland was characterised by medium sized trees, it had low density (86–102 individuals per ha) and low basal area (3.8–5.7 m² per ha). Species richness ranged between 17 and 27 species per ha in the six plots. Only two species were found in all plots, Daniellia oliveri (Caesalpiniaceae) and Prosopis africana (Mimosaceae). Legumes dominated all plots. Wooded grassland and transition woodland had many characteristics of fire-affected vegetation in contrast to woodland. Today wooded grassland encroaches on woodland and transition woodland. Management of the latter two vegetation types should be given priority as they maintain structural and floristic characteristics that are essential to conserve biodiversity and original features of the vegetation, and they are also important for local people who are allowed to make sustainable use of the vegetation.     

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

We examined recruitment patterns in semi-natural pastures and their relationships to disturbance and species composition at different spatial scales (0.01 m² up to 4 m²) of both the vegetation and the seed bank. Possible associations between seedling recruitment and phenology were also studied. The study was performed in four pastures with different management history. Seedling recruitment was generally enhanced by disturbance, with the greatest effect for small-seeded species that germinate in the autumn. The local species-pool contributed to a large extent to seedling recruitment; 83 % of the recruited species were found in the vegetation, whereas 44 % of the recruited species were found in the seed bank. There were a total of 88 species found in the vegetation at different spatial scales; 61 % were recorded at the smallest spatial scale (0.01 m²). The area sampled at this scale comprised 1.25% of the area examined, indicating a small scale structuring of diversity in the pastures. The species number in the vegetation varied between 7 and 14 for the smallest scale (0.01 m²), and between 23 and 42 for the largest scale (4 m²), in the four pastures. The species number in the seed bank was less variable. Like most studies of perennial grasslands, we found no close correspondence between species distribution in the vegetation and in the seed bank. Two of the pastures had a joint management history and exhibited a close similarity of species in the seed bank, despite that one of the pastures was abandoned already in the beginning of this century.     

The cytotoxic and photodynamic inactivation of micro-organisms by Rose Bengal

Rose Bengal was cytotoxic to the following bacteria at the concentrations given in parentheses (highest concentrations of dye in mol/1 at which growth occurred on nutrient medium): Brochothrix thermosphacta and Deinococcus radiodurans (1 times 10^-6 or less); Streptococcus, Micrococcus, Staphylococcus, Bacillus, Arthrobacter and Kurthia spp. (1 times 10^-5–1 x 10^-4), and Pseudomonas spp. and Enterobacteriaceae (5 times 10^-3–1 x 10^-2 or greater). These organisms were killed rapidly when suspended in illuminated (170 μE/m²/s) solutions of Rose Bengal (1 times 10^-4 mol/1) providing oxygen was present. Singlet oxygen was identified as the lethal agent, because the rate of killing was increased by dissolving the dye in deuterium oxide while the organisms were protected against photoinactivation by L-histidine or crocetin. Yeasts from chilled foods were killed in illuminated solutions of Rose Bengal but a light intensity of 315 μE/m²/s was needed for a death rate comparable with that of bacteria. The yeasts present in a range of chilled meat and dairy products failed to form colonies on Rose Bengal (5 times 10^-5 mol/1) media exposed continuously to modest illumination (55–80 μE/m²/s).     

Relative abundance and growth of mussels (Mollusca: Eulamellibranchia) in pools 8,9 and 10 of the Mississippi River

SUMMARY. Twenty-three species of unionid mussels were sampled during the summer of 1975 from a 76-mile reach of the Upper Mississippi River; this represented eight species fewer than were recorded in a 1930–31 survey for the same section of the river. Of the 244 sites sampled in 1975, mussels were found at 116 sites and the ten most productive sites accounted for 40.8% of the 1205 mussels collected. There were as many as thirteen different mussel species taken from one sampling site. The two most abundant species ( Amblema peruviana and Megalonaias gigantea ) constituted 54.5% of the mussels sampled in 1975; these two species made up only 8.3% of the 1930–31 survey. Individuals of the twelve most abundant mussel species were aged, and shell-length growth equations were determined. In addition, shell-free dry weight growth rate and shell weight growth rate was determined for Amblema peruviana . The calculated annual production for A. peruviana (assuming a mean density of 1 per m²) was 482 mg/m² shell-free dry weight, and 14.471 g/m² shell weight.     

Effects of light and temperature on the cell size and some biochemical components in two freshwater cryptophytes

The effects of light and temperature on cell size and cellular composition (chlorophyll, protein, carbohydrate) of two freshwater cryptophytes were studied with batch cultures. Neither of the species had a constant cell size but the size varied with growth conditions. At each temperature the smallest cells were recorded at the lowest experimental photon flux density. The smallest cells of Cryptomonas 979/67 had an average volume of 232 μm³ and the largest ones 1 020 μm³. In Cryptomonas 979/62 the smallest and largest cells measured 4 306 μm³ and 12 450 μm³. Both species increased their cellular chlorophyll content when PFB dropped below 110–120 μmol m^-2 s^-1. The highest and lowest chlorophyll contents of 979/67 were 7.45 fg μm^-3 and 0.55 fg μm^-2 respectively. For 979/62 the corresponding values were 10.23 fg μm^-3 and 0.93 fg μm^-3. In both species the protein content remained stable at PFDs higher than 110–120 μmol m^-2 S^-1. The highest content of protein measured in 979/67 was 638 fg μm^-3 and the lowest 147 fg μm^-3. For 979/62 these values were 1 036 fg μm^-3 and 148 fg μm^-3 respectively. The carbohydrate results were less clear and no pattern either in response to photon flux density or temperature was obvious. The lowest and highest contents recorded for 979/67 were 62 fg μm^-3 and 409 fg μ^-3 and for 979162, 36 fg μm^-3 and 329 fg μm^-3