Spatial and temporal variation in RGR and leaf quality of a clonal riparian plant: Arundo donax

Arundo donax L. is a tall perennial reed classified as an emergent aquatic plant. In California, it has invaded riparian zones, where it acts as a transformer species. Because plant growth and leaf quality influence the effectiveness of management techniques, we sought to determine if these characters varied temporally and spatially in a northern California population of A. donax. Tissue C and N content and C:N ratio varied during the growing season. Leaf N was higher in spring and in plants that were closer to a stream. It was significantly negatively related to the clump's distance from the stream but not related to its elevation relative to the stream. Plants near the stream produced taller stems with more leaves per stem than those more distant from the stream. RGR differed across time and space. It was highest in spring prior to the appearance of flowers on a few stems that were >1 year old within the clumps. Decline in RGR as the growing season progressed coincided with the appearance of branches and flowers on stems <1 year old on a few plants within the studied population. RGR was significantly related to the N content and C:N ratio of leaves on mature stems (>1 year old). This implies that the decrease in stem growth reflected changes in nutrient availability within the entire A. donax clump and not just in the growing stems (<1 year old). These findings have implications for timing of management techniques.     

Biomechanical properties of the emergent aquatic macrophyte Sparganium erectum: Implications for fine sediment retention in low energy rivers

This paper is concerned with the biomechanical properties of the emergent aquatic macrophyte, Sparganium erectum. We present observations of adjustments in the physical characteristics and biomechanical properties of S. erectum during the growing season (April-November) from the River Blackwater, UK. When a pulling device is attached to plant stems to measure their resistance to uprooting, individual plants show remarkable strength in their above- and below-ground biomass (median stem strength when stems break away from the underground biomass, 78 N, median rhizome strength, 39 N) and high resistance to uprooting (median uprooting resistance when entire plants uproot, 114 N). This provides the potential for the species to protect and reinforce the generally soft, silty sediments that it often retains and within which its rhizomes and roots develop in lower energy river environments. There is a propensity for plant stems to break before the plant is uprooted at the beginning and end of the growth season, but for the stems to have sufficient strength in mid season for plant uprooting to dominate. This ensures that rhizome and root systems remain relatively undisturbed at times when the silty sediments in which they grow are poorly protected by above-ground biomass. In contrast, rhizome strength remains comparatively invariant through the growing season, supporting the plant's potential to have a protective/reinforcing effect on fine sediments through the winter when above ground biomass is absent.     

Carbon and nitrogen translocation in response to shading of the seagrass Posidonia sinuosa

Translocation of carbon (C) and nitrogen (N) was investigated in response to shading of the seagrass Posidonia sinuosa in control (ambient light) and shade (below minimum light requirement) treatments after 10 d shading. A mature leaf was incubated in situ in ¹³C- and ¹⁵N-enriched seawater for 2 h and the appearance of the isotopes in the young leaf and adjacent rhizome monitored over 29 d. C and N isotopes gradually reduced in the mature leaf: of ¹⁵N contained in the entire shoot (mature leaf, young leaf and 4 cm rhizome), 95% (control) and 97% (shade) was found in the mature leaf after 2 h incubation and only 75% and 60% remained in the mature leaf after 29 d; 98% and 94% of ¹³C was found in the mature leaf after 2 h, and it had reduced to 36% and 44% after 29 d. This corresponded to an equal increase in the young leaf + rhizome indicating that the mature leaf is a source of these nutrients to the young leaf and rhizome. C translocation from mature leaves was not significantly affected by the shade treatment. In contrast, there was an increase in ¹⁵N taken up by the mature leaves (1.9× higher in the shade), the percent of ¹⁵N translocated to the young leaf and rhizome (24% in control and 40% in shade) and N concentration in the young leaf (1.24% control and 1.41% shade) and rhizome (0.86% control and 0.99% shade). Resorption of C and N was also estimated from changes in the total C and N content of the mature leaf over 29 d. N resorption from the mature leaf contributed up to 63% of young leaf N requirements in the control treatment but only 41% in the shade treatment. We conclude that uptake and translocation of N by mature leaves is a response to shading in P. sinuosa and would provide additional N to growing leaves, enhancing light harvesting efficiency.     

The effects of nitrogen application on growth and nitrogen distribution within the potato canopy

The effect of applying nitrogen (N) fertiliser on the growth and distribution of N within the potato canopy was studied in 1983 and 1984. In both years N was applied either in excess of that required to produce maximum tuber yields, or not at all. The large application of N changed the pattern of canopy growth - stimulating growth of leaves at the top of the stem, particularly lateral branches, for longer during the season, and accelerating the death of (shaded) leaves at the base of the canopy. The pattern of canopy senescence was, therefore, changed from a synchronous to a progressive type. Application of nitrogen fertiliser at supra-optimal rates increased the N contents of leaves, stems and tubers. The extra N in the leaves of these plants was present as reduced N in all leaf positions, and as nitrate (NO^-₃) in the lowermost leaves. In addition, substantial quantities of NO^-₃ were also stored in the stems. Part of this extra N in the canopy was redistributed during subsequent growth, especially to the lateral branches as crop N uptake slowed towards the end of the season. In addition, substantial quantities of N were also potentially available for redistribution to the growing tubers. There was little redistribution of N from the leaves of N-deficient plants. It is suggested that redistribution of N in the canopy of N-replete plants allowed the growth of lateral branches towards the end of the season, thereby maintaining photosynthetically active leaves for longer than N-deficient plants.     

Ambient UV-B radiation reduces PSII performance and net photosynthesis in high Arctic Salix arctica

Ambient ultraviolet-B (UV-B) radiation potentially impacts the photosynthetic performance of high Arctic plants. We conducted an UV-B exclusion experiment in a dwarf shrub heath in NE Greenland (74°N), with open control, filter control, UV-B filtering and UV-AB filtering, all in combination with leaf angle control. Two sites with natural leaf positions had ground angles of 0° (‘level site’) and 45° (‘sloping site’), while at a third site the leaves were fixed in an angle of 45° to homogenize the irradiance dose (‘fixed leaf angle site’). The photosynthetic performance of the leaves was characterized by simultaneous gas exchange and chlorophyll fluorescence measurements and the PSII performance through the growing season was investigated with fluorescence measurements. Leaf harvest towards the end of the growing season was done to determine the specific leaf area and the content of carbon, nitrogen and UV-B absorbing compounds. Compared to a 60% reduced UV-B irradiance, the ambient solar UV-B reduced net photosynthesis in Salix arctica leaves fixed in the 45° position which exposed leaves to maximum natural irradiance. Also a reduced Calvin Cycle capacity was found, i.e. the maximum rate of electron transport (J_max) and the maximum carboxylation rate of Rubisco (V_cmax), and the PSII performance showed a decreased quantum yield and increased energy dissipation. A parallel response pattern and reduced PSII performance at all three sites indicate that these responses take place in all leaves across position in the vegetation. These findings add to the evidence that the ambient solar UV-B currently is a significant stress factor for plants in high Arctic Greenland.     

Hydrothermal and thermal time models for the invasive grass, Arundo donax

Controlled laboratory and field experiments were performed to determine the developmental response to temperature and moisture of Arundo donax, a riparian invasive grass and potential bioenergy crop. A logistic function was parameterized and used to predict thermal times to sprouting and the nine-leaf stage. Consistent estimates of the base temperature (T_b) and base water potential (ψ_b) below which development ceases were obtained from various statistical and mathematical analyses. Estimates of T_b and ψ_b were 12.7 ± 1.7 °C and −1.56 ± 0.43 MPa, respectively, for the median fraction of sprouting rhizomes. Median hydrothermal time to sprouting was 124.1 MPa °Cd under laboratory conditions and median thermal times, or degree-day (°Cd), to sprouting and nine-leaf stage was estimated to be 94 and 129 °Cd under field conditions, respectively. A degree-day is defined as one day (24 h) spent one degree above T_b. Results demonstrated that thermal time alone is sufficient to accurately predict time to sprouting under field conditions. Further, there may be a fixed moisture threshold of about 6% volumetric water content above which sprouting rate was constant. This threshold corresponded very closely to the −1.5 MPa for ψ_b that was estimated under laboratory conditions for the soil typically infested by A. donax. This information is crucial for assessing risk of invasive spread for A. donax.     

Can small water level fluctuations affect the biomass of Nymphaea alba in large lakes?

In this study we investigated above-ground biomass and morphological responses of a floating-leaved plant species, Nymphaea alba, to small spring water level manipulations (0.1–0.5 m) in a large, shallow lake over a 9-year period (1995–2003). A year effect was found in mean annual above-ground plant biomass with higher values found in years of low water levels, 275–339 g DW m⁻² in 1995 and 2003 against 143–198 g DW m⁻² in 1996–2002 (no data transformation). No significant changes in biomass patterns were observed within each season (one summer peak), except in 1995 when a summer decline in biomass occurred. The amplitude and duration of exposure to high water levels affected the spring and annual above ground biomass of N. alba. The plant responded to high spring water levels by producing longer and thinner petioles to preserve leaves from flooding while no significant changes in leaf surface area (except in May) and leaf/petiole biomass ratio were obtained. The results are interpreted with regard to plant adaptations to changing environments (biomass allocation patterns in the different plant organs and stem density) and the effects of other abiotic factors relevant to the size of the system. We concluded that small deviations in spring water level can be driving forces in a large system in controlling the above-ground biomass of this floating-leaved plant.     

Light promotes an increase of cytokinin oxidase/dehydrogenase activity during senescence of barley leaf segments

Following a study of the relationship between cytokinin oxidase/dehydrogenase (CKX) and senescence in darkened barley leaf segments, we have now investigated the influence of light on the in vitro activity of CKX. Seedlings of Hordeum vulgare L. were grown for 8 d under a light/dark regime of 18 h white light and 6 h darkness. Then apical parts of 7 cm length were cut from the first foliage leaves and their bases were placed in water. In segments kept in the dark, the CKX activity measured by cleavage of N₆-(Δ₂-isopentenyl)adenine rose from 0.1 pkat (g FW)⁻¹ to 0.8 pkat (g initial FW)⁻¹ within the first 4 d of incubation. In contrast, in segments kept under the light/dark regime it reached a value of 8.6 pkat (g initial FW)⁻¹ over the same time period. The chlorophyll a content declined slightly slower during light/dark cycling than in darkness. In contrast to segments and isolated laminae, corresponding attached laminae exhibited less CKX activity after 2 d under light/dark conditions than after 2 d in the dark. The activity in attached laminae of first foliage leaves of plants growing in light/dark cycling increased strongly only when the plants were older than 4 weeks. In line with this, the CKX activity in attached laminae of flag leaves of barley growing in fields increased in a late developmental state. The senescence of darkened isolated laminae of Zea mays L. and Phragmites australis (Cav.) Trin. ex Steudel was associated with an enhancement of CKX activity too. Because in most cases a positive correlation between CKX activity and senescence was found, it is likely that the enzyme promotes senescence by destroying cytokinins, which help to keep Poaceae leaves green. Light may promote not only cytokinin degradation but also the formation of bioactive cytokinins in leaf segments.     

Selecting browse plants to supplement cassava peel-based diet for peri-urban small ruminants

Cassava (Manihot esculanta) peel is routinely fed to ruminants in Ghana and most parts of Africa, but the low protein content and lack of suitable protein supplements limits its full exploitation in small ruminant production systems. This study assessed the suitability (degradation characteristics and synchrony between nitrogen release and organic matter degradation) of the leaves of three browse plants, chaya (Cnidoscolus aconitifolius), ficus (Ficus exasperata), and terminalia (Terminalia catappa), as supplements for sheep fed a cassava peel-based diet. Four ruminally fistulated Djallonké wethers were used in a randomized complete block design experiment to determine ruminal disappearance of dry matter (DM), organic matter (OM) and nitrogen (N) from the three leaves and from cassava peels. For chaya, ficus and terminalia, OM content was 93.1%, 83.8% and 90.3% (P < 0.05), respectively, whereas N was 3.4%, 3.1% and 1.7% (P < 0.05, DM basis). The OM and N content in cassava peel was 95.3% and 1.0%, respectively. Ruminal DM disappearance of chaya, ficus, terminalia and cassava peel after 24 h of incubation was 79.0%, 36.4%, 48.0%, and 43.0% (P < 0.05), respectively. The ratio of N released from chaya leaves to OM from cassava peel was 1:16 or lower during 24 h ruminal incubation, compared to 1:31 during the first 8 h and 1:25 after 12 h of incubation of ficus leaves. The ratio for terminalia leaves from 4 to 24 h of incubation ranged between 1:51 and 1:63. The high N content and ideal synchronous release of N and OM ratio of 1:33 (N:OM) made ficus leaves the most suitable supplement among the three browse leaves, with the potential to supply adequate N to optimize ruminal microbial protein synthesis in sheep on a cassava peel-based diet.     

Seasonal growth of tree lupins (Lupinus arboreus) and the effect of defoliation on leaf production

The growth of tree lupins was investigated in two experiments. In the first, two ages of plant, 4-wk-old seedlings and 1-year-old plants, were transplanted into a ryegrass sward in an upland environment. Growth, in terms of leaf production, branching and stem elongation, was measured over two successive growing seasons. Plant dry matter and nutrient contents were determined at the beginning and end of each growing season. In the first summer, the rate of production of new leaves on the main stem of seedling plants averaged 1.8 leaves per wk and main stem length increased from 5 to 67 cm. On older plants, where floral apices had been initiated on main and primary stems, there was a 3–10 fold increase in secondary branch length. In the second season, there was no effect of plant age on rates of leaf appearance or stem extension; dry matter production was higher than in the first season. In the second experiment, the effect of removal of 0%, 50% or 100% of fully expanded leaves on the subsequent growth of 23-wk-old plants was investigated. During the 7-wk growth period, defoliation promoted the rate of production of mature leaves, and area and dry weight of new laminae were slightly higher in defoliated plants. Defoliation did not affect the concentrations of N, P or K in the new laminae, but P and K concentrations in petioles of defoliated plants were significantly higher than those in intact plants. The results from the experiments are discussed in relation to the potential use of tree lupins as nurse species and biomass crops in hill and upland environments of the UK.     

Germination and early growth of Nymphaea odorata at different water depths

We experimentally determined the effects of water depth on seed germination and seedling growth and morphology, and we documented the transition from submerged to emergent plants in the white water lily, Nymphaea odorata. Seeds of N. odorata were germinated at 30, 60, and 90 cm water depth in outdoor mesocosms and percent germination and morphology measured after a month. The presence of self-seeded seedlings in pots at the same 3 water levels was also recorded over two years. To examine juvenile growth, seeds planted in soil were placed at the same mesocosm depths; germination and growth were monitored for three months, when the plants were harvested for morphological and biomass measurements. N. odorata germinated equally well in 30, 60 and 90 cm water; seedlings grew as submerged aquatics. After one month, seedlings in 90 cm water had less biomass than those in 30 cm (1.1 vs. 3.3 mg and 1.0 vs. 1.8 mg for different seed sources, respectively) and allocated relatively more biomass to shoots (97.5 vs. 67.8% and 73.1 vs. 58.0%, respectively). Seedlings in 60 cm water were intermediate. After 3 months of submerged growth, plant biomass remained less in 90 vs. 60 and 30 cm water (22.5 vs. 36.4 and 33.3 mg, respectively). Plants in 90 and 60 cm water had greater biomass allocation to shoots than plants in 30 cm water (85.7 and 72.6% vs. 64.4%, respectively) and produced larger laminae on longer petioles (lamina length = 33.3 vs. 25.2 mm in 90 vs. 30 cm; petiole length = 99.0 vs. 36.0 mm, respectively). After about 3 months, submerged plants produced floating leaves that had 39% shorter laminae but 267% to 1988% longer petioles than submerged leaves on the same plant. Lamina length to width allometric relations of submerged leaves were >1 at all water levels, distinguishing them from the equal allometry of adult floating leaves. The switch from production of submerged to emergent leaves resembles submergence-escape growth in other aquatics, but because the seedlings have been submerged throughout their life, submergence itself cannot be the stimulus to produce emergent leaves in these totally immersed plants. Our data show that N. odorata plants can establish from seeds in up to 90 cm water and that seedlings grow as submerged aquatics until they switch abruptly to production of floating leaves.     

Contrasting responses of seagrass transplants (Posidonia australis) to nitrogen, phosphorus and iron addition in an estuary and a coastal embayment

Addition of nutrients to sediments has been proposed as a means of enhancing transplantation success in seagrasses. The effects of nutrient and iron additions to natural sediments on the growth and morphology of Posidonia australis transplants were evaluated in underwater plots in two contrasting environments: a coastal embayment (Princess Royal Harbour) with sandy sediments and little riverine input, and an estuary (Oyster Harbour) with organic-rich sediments and subject to seasonal river flow from a large rural catchment. Sixty six planting units spaced 1 m apart were transplanted in situ in each location. Nitrogen (N) and phosphorus (P) were added in a randomized factorial design using slow release fertilizer granules at the start of the experiment and repeated every 4-5 months for 2 years. In a concurrent experiment, chelated iron Fe EDTA was added to modify the sediment sulphur cycle.In Oyster Harbour, the addition of N significantly increased leaf N concentrations but reduced total biomass and biomass of leaves. Addition of P significantly increased leaf P concentrations and number of living leaves per transplant, leaf area, leaf length, length of longest rhizome axis and total rhizome length. Combined N + P addition resulted in a significant increase in leaf P concentrations and leaf area per plant only. In Princess Royal Harbour, addition of N produced significant increases in leaf variables (total and leaf biomass, number of shoots and living leaves, leaf area, and leaf length) but there were no significant differences observed in below ground plant parts (rhizomes). Addition of P had no significant effects on any growth measurements. Addition of N + P combined increased number of living leaves and leaf area significantly. δ¹⁵N in mature leaf tissue were significantly more negative for N and N + P treatments at both locations.Our results indicated that N limitation was occurring in the coastal embayment, Princess Royal Harbour whereas in the more estuarine Oyster Harbour, P was limiting plant growth. Addition of FeEDTA produced equivocal results at both sites and we suggest these results are confounded by the addition of N and C in the EDTA. We caution the use of nutrient addition to transplants of slow growing seagrasses such as P. australis without a thorough understanding of the nutrient status of the system, estuarine or coastal embayment, in which they are to be transplanted.     

Morphological and physiological adaptive traits of Mediterranean narrow endemic plants: The case of Centaurea gymnocarpa (Capraia Island,Italy)

A case study on Centaurea gymnocarpa Moris & De Not., a narrow endemic species, was carried out by analyzing its morphological, anatomical, and physiological traits in response to natural habitat stress factors under Mediterranean climate conditions. The results underline that the species is particularly adapted to the environment where it naturally grows. At the plant level, the above-ground/below-ground dry mass (1.73 ± 0.60) shows its investment predominately in the above-ground structure with a resulting total leaf area per plant of 1399 ± 94 cm². The senescent attached leaves at the base of the plant contribute to limit leaf transpiration by shading soil around the plant. Moreover, the dense C. gymnocarpa leaf pubescence, leaf rolling, the relatively high leaf mass area (LMA = 12.3 ± 1.3 mg cm⁻²) and leaf tissue density (LTD = 427 ± 44 mg cm⁻³) contribute to limit leaf transpiration, also postponing leaf death under dry conditions. At the physiological level, a relatively low respiration/photosynthesis ratio (R/P_N) in spring results from high R [2.26 ± 0.59 μmol (CO₂) m⁻² s⁻¹] and P_N [12.3 ± 1.5 μmol (CO₂) m⁻² s⁻¹]. The high photosynthetic nitrogen use efficiency [PNUE = 15.5 ± 0.4 μmol (CO₂) g⁻¹ (N) s⁻¹] shows the large amount of nitrogen (N) invested in the photosynthetic machinery of new leaves, associated to a high chlorophyll content (Chl = 35 ± 5 SPAD units). On the contrary, the highest R/P_N ratio (1.75 ± 0.19) in summer is due to a significant P_N decrease and increase of R in response to drought. The low PNUE [1.5 ± 0.2 μmol (CO₂) g⁻¹ (N) s⁻¹] in this season is indicative of a greater N investment in leaf cell walls which may contribute to limit transpiration. On the contrary, the low R/P_N ratio (0.05 ± 0.02) in winter is resulting from the limited enzyme activity of the respiratory apparatus [R = 0.23 ± 0.08 μmol (CO₂) m⁻² s⁻¹] while the low PNUE [3.5 ± 0.2 μmol (CO₂) g⁻¹ (N) s⁻¹] suggests that low temperatures additionally limit plant production. The experiment of the imposed water stress confirms that the C. gymnocarpa growth capability is in conformity with the severe conditions of its natural habitat, likewise as it may be the case with others narrow endemic species that have occupied niches with similar extreme conditions.     

Alternanthera bettzickiana (Regel) G. Nicholson,a potential halophytic ornamental plant: Growth and physiological adaptations

Exploration and cultivation of salt tolerant plants is a very effective strategy for utilization of salt affected soils. In this investigation, physiological traits that are conducive for salt tolerance of the ornamental plant Alternanthera bettzickiana, Amaranthaceae, were explored. A. bettzickiana was grown on soil substrate having six salinity levels (2.86, 10, 20, 30, 40 and 50 dS m⁻¹). It was observed that this plant can grow even at a salinity level of 40 dS m⁻¹. The survival rate of this plant was 75, 42 and 0% at salinity levels of 30, 40 and 50 dS m⁻¹, respectively. A. bettzickiana plants produced 30.3% less biomass than controls at the salinity level of 20 dS m⁻¹ and even less under still higher salt stress. Photosynthesis continued even at the salinity level of 40 dS m⁻¹, though its rate was reduced to 59% in plants exposed to such salinity relative to plants not affected by salinity. Total soluble proteins values in leaf and stem showed a gradual increase when plants were exposed to increasing salt stress. Plants growing at the high salinity level showed highest decrease in leaf nitrate reductase activity. A. bettzickiana plants accumulated less Na⁺ in shoot as compared to root when grown under salt stress. It can be characterized as a salt-tolerant glycophyte that could be used for greening of salt affected soils.     

Remobilization patterns of C and N in soybeans with different sink-source ratios induced by various night temperatures

The effects of increased sink-source ratios, induced by elevating night temperatures, on remobilization of ¹⁴C-assimilates and N within field-grown soybeans (Glycine max [L.] Merr.) was investigated from preflowering to maturity. Raising the mean minimum night temperature for the entire growing season from 10 (check, uncontrolled) to 16°C increased seed growth without appreciable effect on final leaf area. Increasing this temperature to 24°C increased seed growth and reduced final leaf area. Leaves, stems, petioles, and pods acted as intermediate storage sites for ¹⁴C assimilates. Only plants with higher night temperatures remobilized some of the stored assimilates during the period of rapid seed growth. Even the seeds in the 24°C plants with the largest sink-source ratios did not utilize all the C-assimilates potentially available for remobilization. Nitrogen was readily remobilized from petioles, stems, and pods of all treatments as early as the beginning of seed development, but from the leaves only during late seed-filling. However, only plants with elevated night temperatures tended to remobilize all of the available N from vegetative tissues and pods. We concluded that a larger portion of stored assimilates may be remobilized to the seed if a strong seed sink can be sustained. It also appeared that with increasing sink-source ratios, N shortage might limit seed yield before a lack of C-assimilates would. A proposed model for soybean assimilate demand, distribution, partitioning, and remobilization is presented.     

Effect of cadmium on growth and micronutrient distribution in wild garlic (Tulbaghia violacea)

Tulbaghia violacea Harv. (Alliaceae) is one of the few medicinal plants that is also frequently used as a leafy vegetable. Application of cadmium (Cd) at 2 and 5 mg/L to T. violacea plants of various sizes (small 8–10 g, medium 16–20 g, large 80–95 g) elicited a difference in growth response, Cd accumulation and micronutrient distribution. Application of Cd up to 5 mg/L had no significant effect on growth parameters of large-sized plants while leaf length and fresh weight of leaves of the medium-sized plants decreased with application of Cd at 2 mg/L, and 5 mg/L. Cadmium significantly decreased the number of leaves in small-sized plants. Small plants accumulated more Cd in the leaves than medium or large-sized plants. Application of Cd at 2 and 5 mg/L lowered the leaf Cu, Fe, Mo and Zn contents in small and medium-sized plants but had no effect on the micronutrients in large-sized plants. This study indicates that T. violacea has the ability to accumulate Cd. In addition, plant size plays an important role with regards to Cd accumulation and elemental distribution. The results presented in this study include the first report on the nutritional status of T. violacea leaves.     

Differential effects of nitrogen and phosphorus enrichment on growth of dwarf Avicennia marina mangroves

The effects of N and P enrichment were investigated on growth and physiological responses of dwarf Avicennia marina mangroves in a hypersaline (58 ± 8 psu) field site in Richards Bay, South Africa. It was hypothesized that at high salinities mangroves allocate more resources to roots than shoots, and that nutrient enrichment with N and P will shift resource allocation to shoots and enhance growth and productivity. In unvegetated areas of the dwarf zone, 1-year-old A. marina seedlings were planted in pots and enriched bimonthly with N, P, N + P, or remained unfertilized (control-C), and growth and morphology of plants were monitored for 2 years. Enrichment with N and N + P shifted resource allocation to shoots from 38% to 55%, and increased dry biomass accumulation by over 500%, compared to the control treatment. In the N and N + P treatments, plant height, number of leaves, leaf chlorophyll content and photosynthesis increased by over 50%, 330%, 30% and 30%, respectively, compared to the C and P treatments. Enrichment with N and N + P increased N concentrations in roots by over 60% (from 1.0 ± 0.1% to 1.6 ± 0.2% of dry mass) and in shoots by over 100% (from 1.3 ± 0.1% to 2.7 ± 02% of dry mass). Plants enriched with P alone were similar to those of the control. This study has demonstrated that dwarf A. marina in Richards Bay is N limited, and that N enrichment shifts resource allocation from roots to shoots and increases growth and productivity.     

Evaluation of the transpiration rate of lotus using the stem heat-balance method

To reveal the mechanism of transpiration by hydrophytes in the field, it is necessary to evaluate the transpiration rate without the effect of the evaporation from the water surface. In order to test the suitability for evaluating the transpiration rate of lotus (Nelumbo nucifera Gaertn.) leaves in the field, stem heat-balance method was applied and the obtained sap-flow rate was compared with the transpiration rate measured by weighing and with the overall canopy evapotranspiration rate by means of the eddy covariance technique. The transpiration rate estimated with the sap-flow measurements showed good agreement with that obtained from the weighing method. Lotus has many air canals in its petiole to carry oxygen-rich air to the rhizome and methane- and carbon dioxide-rich air back to the atmosphere, but there was little effect of the mass flow of air through these canals on the sap-flow rates. In the field observations, the canopy evapotranspiration rate (0.28 mm h⁻¹ at maximum) was nearly equal to the sum of the transpiration rate from all sunlit leaves (0.30 mm h⁻¹), and the contribution of the transpiration from shaded leaves and evaporation from the water surface was considered to be minor in the seasons when the leaves were fully developed. Evaluation of bulk leaf conductance revealed that the conductance in the leaf boundary layer of lotus could be low (ca. 0.23 mol m⁻² s⁻¹) because of its large leaf area. The low conductance in the leaf boundary layer would increase leaf temperature, which, in turn, would generate air circulation within the plant's ventilation system. Because there was a linear relationship between transpiration rate and the leaf-to-air vapor-pressure deficit, with no apparent maximum, high vapor-pressure deficits (3.4 kPa at maximum) did not appear to cause significant stomatal closure in lotus plants. The stomata of lotus leaves play a role as air inlets to carry oxygen-rich air to the rhizome, so their low sensitivity would help to increase air intake.     

Do freshwater plants have adaptive responses to typhoon-impacted regimes?

In tropical regimes, cyclones exert great influence on the local aquatic habitats. The objective of our study was to investigate if aquatic plants have an adaptive response to typhoon influence. Population traits of six aquatic species in different life-forms (emergent species: Scirpus triangulatus, Eleocharis plantagineiformis, Rotala rotundifolia, Eriocaulon buergerianum; submerged: Blyxa echinosperma; floating-leaved: Nymphoides indica) were investigated to compare intraspecific variations in high and low typhoon-impacted regions on Hainan Island in southern China. In the high typhoon-impacted region, there was greater belowground biomass allocation in both emergent and floating-leaved species. The ratio of belowground to total biomass of each emergent was 41% (P = 0.028), 38% (P = 0.034), 27% (P = 0.040), 19% (P = 0.043) greater respectively, and floating-leaved N. indica was 40% (P = 0.014) greater than in the low typhoon region. The stem height of relatively tall emergent species (S. triangulatus and E. plantagineiformis) was 35% (P = 0.033), 42% (P = 0.046) lower, and floating-leaved species N. indica had decreased leaf area (49%, P < 0.001) and number (30%, P < 0.001) on water surface in the high typhoon-impacted region than in the low. These adaptations of the plants will reduce their risk of mechanical damage from strong winds or wind-induced currents. Submerged species in the study showed no variation in traits between the high and low typhoon-impacted regions.     

Seasonal growth, δ13C in leaves and stem,and phloem structure of birch (Betula pendula) under low ozone concentrations

Summary The growth of potted birch cuttings (one clone of Betula pendula) was studied under low O₃ concentrations (0, 0.050, 0.075, 0.100 l l^-1) throughout an entire growing season. With increasing O₃ dose, 20–50% of all leaves formed were prematurely shed, while 40–70% of the remaining foliage displayed advanced discoloration by the end of the season. Ozonation affected the S, P and N concentration of leaves and increased ¹³C in leaves and stem, while the CO₂ assimilation rate declined with increasing CO₂ concentration in mesophyll intercellulars. While whole-plant production correlated negatively with the O₃ dose, ozone increased the specific leaf weight (i.e. leaf weight/leaf area, SLW) but decreased the ratios of stem weight/stem length and root/shoot biomass. Neither the latter ratio nor SLW changed in experimentally defoliated control plants, whereas in ozonated plants starch accumulated along leaf veins and phloem tissue was deformed in the leaf petioles and the stem. Only in early summer was the relative growth rate higher in the ozonated than in the control plants. The ratio of whole-plant biomass production versus total foliage area formed was lowered under O₃ stress. However, when relating biomass to the actual foliage area present due to leaf loss, this ratio did not differ between treatments. Similarly the ratio of actual foliage area versus basal stem area in cross-section did not differ. Overall, whole-plant production was strongly determined by O₃-caused changes in crown structure and began to be limited at O₃ doses (approximately 180 l l^-1 h) similar to those of rural sites in Central Europe.