Clonal integration enhances flood tolerance of Spartina alterniflora daughter ramets

Recently, considerable attention has been paid to the invasion of the clonal plant Spartina alterniflora into coastal wetlands at lower elevations. In this experiment, we tested whether clonal integration improved flood tolerance in S. alterniflora daughter ramets. Daughter ramets at two growth stages (young and old ramets) were flooded to water levels of 0, 9 and 18 cm above the soil surface, and the rhizomes between mother and daughter ramets were either severed or left intact. Biomasses of connected ramets grown in controls or in shallow and deep water treatments were 119%, 108% and 149% higher in the old ramet group than those of severed ramets, respectively, whereas they were 3.0, 3.3 and 11.2 times higher in the young ramet group, respectively. At the end of the experimental period, the shoot height, connected with young ramets, in shallow and deep water treatments increased by 19% and 26%, respectively, over that in the control treatments, whereas the old ramets increased by 11% and 39%, respectively. In contrast, the shoot height of the severed young ramets was 27% and 26% lower in shallow and deep water treatments than in the control treatment, respectively. However, the shoot height of the severed old ramets remained constant with increasing water depth. We conclude that clonal integration enhances the flood tolerance of S. alterniflora daughter ramets, and the trait of clonal integration plays more important roles in severe flooding stress conditions and at early growth stages.     

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.     

Differences in the structure, growth and survival of Parasenecio yatabei ramets with contrasting water relations on the slope of a stream bank

Parasenecio yatabei (Asteraceae), a summer-green perennial herb, is widely distributed on sloping mountain stream banks in cool-temperate zone forests of Japan. We investigated the growth pattern, leaf longevity and leaf water relations of vegetatively independent plants (ramets) growing in two contrasting soil water conditions, that is, upper and lower stream banks (U ramets and L ramets, respectively). The objective of the present study was to clarify the physiological and morphological responses of the ramets to soil water conditions. Dry matter allocation to subterranean parts was higher in U ramets than in L ramets. The U ramet leaves survived for approximately 2 months longer than L ramet leaves. The ratio of subterranean part to aerial part dry matter was greater in U ramets than L ramets. Leaf mass per leaf area (LMA) tended to be greater in U ramets than L ramets throughout the growing season. The leaf bulk modulus of elasticity at full hydration was significantly higher in U ramets. Thus, ramet growth patterns and morphological traits varied with changing soil water conditions. The greater longevity of U ramet leaves may play a role in compensating for the reduced annual net carbon gain caused by lower photosynthetic activity. U ramets growing in environments with less water availability achieved high water-use efficiency by a high passive water absorption capacity via a progressed root system and high productivity via longer leaf longevity with higher LMA and elasticity. Therefore, P. yatabei growing along mountain streams could have the ability to colonize the upper bank through higher survivorship based on these traits.     

Response of physiological integration in Trifolium repens to heterogeneity of UV-B radiation

Physiological integration has been documented in many clonal plants growing under resource heterogeneity. Little is still known about the response of physiological integration to heterogeneous ultraviolet-B radiation. In this paper, the changes in intensity of physiological integration and of physiological parameters under homogeneous and heterogeneous ultraviolet-B radiation (280-315 nm) were measured in order to test the hypothesis that in addition to resource integration a defensive integration in Trifolium repens might exist as well. For this purpose, homogeneous and heterogeneous ultraviolet-B radiation was applied to pairs of connected and severed ramets of the stoloniferous herb Trifolium repens. Changes in intensity of water and nutrient integration were followed with acid fuchsin dye and ¹⁵N-isotope labeling of the xylem water transport. In order to assess the patterns of physiological integration contents of chlorophyll, ultraviolet-B absorbing compounds, soluble sugar and protein were determined and activities of superoxide dismutase (SOD) and peroxidase (POD) measured. When ramets were connected and exposed to heterogeneous UV-B radiation, the velocity of water transportation from the UV-B treated ramet to its connected sister ramet was markedly lower and the percentage of ¹⁵N left in labelled ramets that suffered from enhanced UV-B radiation was higher and their transfer to unlabelled ramets lower. In comparison with clones under homogeneous ultraviolet-B radiation, the content of chlorophyll, ultraviolet-B absorbing compounds, soluble sugar and activities of SOD and POD increased notably if ultraviolet-B stressed ramets were connected to untreated ramets. Chlorophyll and UV-B absorbing compounds were shared between connected ramets under heterogeneous UV-B radiation. This indicated that physiological connection improved the performance of whole clonal plants under heterogeneous ultraviolet-B radiation. The intensity of physiological integration of T. repens for resources decreased under heterogeneous ultraviolet-B radiation in favor of the stressed ramets. Ultraviolet-B stressed ramets benefited from unstressed ramets by physiological integration, supporting the hypothesis that clonal plants are able to optimize the efficiency of their resistance maintaining their presence also in less favorable sites. The results could be helpful for further understanding of the function of heterogeneous UV-B radiation on growth regulation and microevolution in clonal plants.     

Estimating Arundo donax ramet recruitment using degree-day based equations

Arundo donax is a tall perennial reed. Once established, it spreads by producing new shoots (ramets) from rhizomes. We performed two separate experiments to test the hypotheses that temperature (7, 8, 14, 16, or 20 °C) and combinations of temperature and nitrate concentration (0, 0.3, 0.6,1.2, 2.4, 3.6, 4.8, and 6.0 mg/l nitrate) regulated the initiation of ramet production. No ramets emerged from rhizome sections at 7 or 8 °C, but ramets emerged at 14, 16, and 20 °C. Neither time to ramet emergence nor the number of ramets that emerged was influenced by nitrate level in the watering solution. We used the above results in combination with ramet emergence data from plants grown outdoors at Davis, California to develop degree-day equations for three separate ramet cohorts. When compared to ramet emergence from different plants in different years, there was very good agreement between predicted and actual ramet emergence indicating that these equations provide a realistic representation of processes involved in ramet emergence. This is an important step in developing integrated management plans for this species.     

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.     

Anatomical and nutritional requirements for induction and sustained growth in vitro of Cymodocea nodosa (Ucria) Ascherson

In vitro methods of plant micro-propagation are being considered as a possible solution for the decline in seagrass communities registered worldwide. To achieve successful plant micro-propagation, culture conditions are commonly adjusted empirically within almost species-specific conditions, to comply to the following three conditions: (i) culture establishment (ii) shoot production and (iii) rooting and hardening for planting in soil. Cultures of Cymodocea nodosa were established from axenic explants of the apical meristem (approx. 0.5 cm) which regenerated new leaf or produced leaf regenerating calli (5% of cultivated explants) in media containing 10⁻⁶ M of the cytokinin analogue TDZ. Longer ramet explants, not fully axenic, containing internode with leaf and roots were also affected by 10⁻⁶ M cytokinins and auxin type of regulators, as they promoted leaf extension (in cm), particularly GA. None of the explants progressed further to massive shoot propagation and new plantlet production. Instead, experiments made with ramet explants which simulated potential produced plantlet revealed that there seems to be a strong interaction within leaf, rhizome and root, since the carbon fixed in the leaf was rapidly translocated to the rest of the tissue (50% in the roots in a FW basis). The explants preferred ammonium and dihydrogen inorganic phosphate as a nutrient source, efficiently assimilating the former regardless of whether such were added to the underground or surface tissue. However, underground tissue was required to maintain P status in the cultivated explants.     

Distylic Hottonia palustris shows high reproductive success in small populations despite low genetic variability

Hottonia palustris L. (Primulaceae) is characterized by a heteromorphic incompatibility system. The strategy of distylic ramets is believed to promote outcrossing, to maintain overall genetic diversity and to prevent inbreeding depression. In spite of this distyly, an extremely low amount of allozymic diversity was observed in 545 individual ramets from 14 populations in Flanders (Belgium). A possible explanation for such low genetic variation is discussed in relation to the vegetative propagating abilities and the ecological niche width of the species. In contrast to the uniformity in allozymes as well as to the feature of single morph populations, there was a high variability in reproductive success between populations such as the number of seeds per ramet (425–2633), the number of flowers per ramet (9–36) and the mean weight of seeds (0.03–0.17 mg). Small populations and even those consisting of only one style morph may show a high reproductive success. As a whole, H. palustris showed a negative relationship of reproductive success with the surface area of its populations.     

Leaf recruitment and elongation: an adaptive response to flooding in Villarsia reniformis

Villarsia reniformis (Menyanthaceae) responds to flooding by rapid leaf elongation and continual recruitment of young, submerged leaves (4.3–6.5 per week). Leaf production is influenced by nutrient availability and water depth. Leaves are submerged and die as the water level rises, but are replaced by younger leaves able to broach the surface. Young petioles may elongate at more than 10 cm per day, but lose the ability to elongate after the blades are exposed to air more than twice. Young petioles produce new cells and existing cells elongate, but in older petioles fewer new cells are produced and cell elongation, whilst limited, is the main mechanism for petiole elongation. Continual recruitment implies a high cost for production of structural tissue, but ensures that leaves capable of rapid extension are within reach of the water surface and the plants can respond quickly to flooding.     

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.     

Shifting effects of physiological integration on performance of a clonal plant during submergence and de-submergence

Background and Aims

Submergence and de-submergence are common phenomena encountered by riparian plants due to water level fluctuations, but little is known about the role of physiological integration in clonal plants (resource sharing between interconnected ramets) in their adaptation to such events. Using Alternanthera philoxeroides (alligator weed) as an example, this study tested the hypotheses that physiological integration will improve growth and photosynthetic capacity of submerged ramets during submergence and will promote their recovery following de-submergence.

Methods

Connected clones of A. philoxeroides, each consisting of two ramet systems and a stolon internode connecting them, were grown under control (both ramet systems untreated), half-submerged (one ramet system submerged and the other not submerged), fully submerged (both ramet systems submerged), half-shaded (one ramet system shaded and the other not shaded) and full-shaded (both ramet systems shaded) conditions for 30 d and then de-submerged/de-shaded for 20 d. The submerged plants were also shaded to very low light intensities, mimicking typical conditions in turbid floodwater.

Key Results

After 30 d of submergence, connections between submerged and non-submerged ramets significantly increased growth and carbohydrate accumulation of the submerged ramets, but decreased the growth of the non-submerged ramets. After 20 d of de-submergence, connections did not significantly affect the growth of either de-submerged or non-submerged ramets, but de-submerged ramets had high soluble sugar concentrations, suggesting high metabolic activities. The shift from significant effects of integration on both submerged and non-submerged ramets during the submergence period to little effect during the de-submergence period was due to the quick recovery of growth and photosynthesis. The effects of physiological integration were not found to be any stronger under submergence/de-submergence than under shading/de-shading.

Conclusions

The results indicate that it is not just the beneficial effects of physiological integration that are crucial to the survival of riparian clonal plants during periods of submergence, but also the ability to recover growth and photosynthesis rapidly after de-submergence, which thus allows them to spread.     

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.     

Effects of water depth and seed provenance on the growth of wild rice (Zizania aquatica L.)

Annual wild rice (Zizania aquatica L.), a species of conservation concern, is an ecologically and culturally important aquatic grass found in stands in the near shore habitats of lakes and rivers in the Midwest and along the eastern coast of North America. This study examined the effects of water depth and seed provenance on the early growth of three Indiana wild rice stands (collected from two lakes) under greenhouse conditions in 2009. Plants were grown at water depths of 46 cm, 23 cm, 0 cm, or −15 cm and harvested either at the first floating leaf stage or at 48 days after transplanting. Wild rice growth was affected by both water depth and seed provenance. The dry weight of roots, stems, leaves, and inflorescences, total biomass, number of tillers, number of leaves, and total leaf area were the lowest in the −15 cm treatment. These vegetative growth parameters also decreased with increasing water depth from the 0 cm treatment. Differences in growth between seed sources were found, supporting the hypothesis that genetic differences among relatively isolated wild rice stands may influence the success of efforts to conserve this species.     

Competitive Performance of Nymphoides Peltata (Gmel.) O. Kuntze Growing in Microcosm

Two experiments were conducted to investigate the effects of competition on growth and performance of Nymphoides peltata (Gmel.) O. Kuntze in microcosm. Part of the research on growth and biomass allocation of N. peltata in response to competition had been reported early (Wu, Z. & D. Yu, 2004, Hydrobiologia 527: 241–250). This paper focuses on the morphological variations of N. peltata under competitive pressure. First, competition between N. peltata and Zizania latifolia (Griseb.) Turcz. ex Stapf. was assigned with the densities of N. peltata to Z. latifolia ratios of 4:0, 4:2, 4:4 and 4:8. Water surface coverage, surface area per leaf blade and number of leaves per plant of N. peltata all declined significantly with increasing density of competitor. Similar results were also found for petiole length and density of branching. However, the variations of planting density did not significantly affect the number of ramets per plant and the stolon length of N. peltata. Second, competitions between N. peltata and emerged Z. latifolia, floating-leaved Trapa bispinosa Roxb. and submerged Myriophyllum spicatum L. were also studied simultaneously. The results showed that significant difference was only found for the water surface coverage of N. peltata. No other significant differences were found for the number of ramets per plant, number of leaves per plant, density of branching, surface area per leaf blade, petiole length, and stolon length of N. peltata. Our studies indicate that N. peltata presents morphological variations when it is growing with Z. latifolia, such that the growth of above-ground parts decrease (i.e., leaf number, petiole length, branching density) and the growth of below-ground parts remains stable (i.e., stolons length). However, N. peltata does not show apparent differences in morphology when it is growing with T. bispinosa or M. spicatum. Accordingly, we conclude that the growth of N. peltata may be apparently inhibited by the presence of Z. latifolia, while T. bispinosa and M. spicatum may have little impact on the growth and performance of N. peltata.     

Increased nutrient supply facilitates acclimation to high-water level in the marsh plant Deyeuxia angustifolia: The response of root morphology

Both water level and nutrient availability are important factors influencing the growth of wetland plants. Increased nutrient supply might counteract the negative effects of flooding on the growth of the fast-growing species. Experimental evidence is scarce and the mechanism is far from clear. The aim of this study is to identify the role of nutrient availability in acclimation to high-water level by investigating the growth and root morphology of the marsh plant Deyeuxia angustifolia, one of the dominant species in the Sanjiang Plain, China. Experimental treatments included two water levels (0 and 10 cm, relative to soil surface) and three levels of nutrient supply (0, 0.5 and 1 g fertilizer per container). High-water level usually led to decreased biomass accumulation, shoot mass and root mass, whereas biomass accumulation was unaffected by water level at the highest nutrient level, indicating that high-nutrient availability played a role in compensating for the growth loss induced by the high-water level. Increased nutrient supply led to decreased root length in 0 cm water-level treatments, but root length increased with nutrient supply in the 10 cm water-level treatments. High-water level usually led to a lower lateral root density, lateral root:main root length ratio and the diameter of main roots and laterals, whereas increased nutrient supply resulted in thicker main roots or laterals, and a higher total root length, lateral root density and lateral root:main root length ratio. These data indicate that the growth of D. angustifolia is restrained by high-water level, and that increased nutrient supply not only ameliorates root characteristics to acclimate to high-water level but also results in a high-total root length to facilitate nutrient acquisition.     

How internode length,position and presence of leaves affect survival and growth of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> after fragmentation?

Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in stoloniferous plants. After fragmentation storage in stolon internodes and leaves may enhance survival and growth of stoloniferous plants. We hypothesize that (1) increasing length of the internode attached to the ramet and (2) presence of leaves will increase ramet survival and growth, and that (3) internode positions (before or after the ramet or both) will also play a role. We tested these hypotheses with the stoloniferous, invasive herb Alternanthera philoxeroides. In one experiment, we measured survival and growth of the ramets either without stolon internode (0 cm in length) or attached with internodes of 2, 4, 6 and 8 cm and either with or without leaves. In the other experiment, we measured survival and growth of the ramets attached with a proximal internode (before the ramet), a distal internode (after the ramet) or both. Increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower. These results support the hypotheses and suggest that storage in stolons and leaves may be of great significance for clonal plants in frequently disturbed habitats and may contribute greatly to the invasiveness of A. philoxeroides.     

Clonal integration improves compensatory growth in heavily grazed ramet populations of two inland-dune grasses

Herbaceous species possess several mechanisms to compensate for tissue loss. For clonal herbaceous species, clonal integration may be an additional mechanism. This may especially hold true when tissue loss is very high, because other compensatory mechanisms may be insufficient. On inland dunes in northern China, we subjected Bromus ircutensis and Psammochloa villosa ramets within 0.5 m×0.5 m plots to three clipping treatments, i.e., no clipping, moderate (50% shoot removal) and heavy clipping (90% shoot removal), and kept rhizomes at the plot edges connected or disconnected. Moderate clipping did not reduce ramet, leaf or biomass density of either species. Under moderate clipping, rhizome connection significantly improved the performance of Psammochloa, but not that of Bromus. Heavy clipping reduced ramet, leaf and biomass density in the disconnected plots of both species, but such negative effects were negated or greatly ameliorated when the rhizomes were connected. Therefore, clonal integration contributed greatly to the compensatory growth of both species. The results suggest that clonal integration is an additional compensatory mechanism for clonal plants and may be important for their long-term persistence in the heavily grazed regions in northern China.     

N:P ratios, δN fractionation and nutrient resorption along a nitrogen to phosphorus limitation gradient in an oligotrophic wetland complex

The vegetation N:P ratio is thought to be a diagnostic indicator of the nature of nutrient limitation in wetland vegetation. It should therefore be closely linked to other indicators of nutrient acquisition and conservation, such as nitrogen stable isotope fractionation (δ¹⁵N), nutrient resorption efficiency (RE) and resorption proficiency (RP). However, the interrelationships among these traits and the N:P ratio remain unclear. We compared tissue nutrient concentrations, N:P ratios, δ¹⁵N fractionation, RE, and RP along an N to P limitation gradient in an oligotrophic wetland valley in the South Island of New Zealand. Within the valley, the soil TN:TP ratio increased from 1.3 to 18.0 in three discrete wetlands along the gradient. In pooled data from all vegetation communities within each site, the mass-based vegetation N:P ratio correlated significantly (r² = 0.35, P < 0.01) to soil TN:TP ratios and increased from 10.2 ± 2.7 to 13.5 ± 3.6 along the N to P limitation gradient. This was accompanied by an increase in tissue δ¹⁵N enrichment from 2.05 ± 1.12‰ to 6.27 ± 1.70‰, consistent with more open N cycling and lower N demand. These trends held within all vegetation types, but were particularly strong in a Typha orientalis (C-strategist) community (soil TN:TP vs vegetation N:P correlation r² = 0.78, P < 0.001; δ¹⁵N increase from 1.81 ± 0.44‰ to 7.73 ± 1.79‰). The individual N and P concentrations and retention patterns were more species-specific and less responsive to the nutrient limitation gradient. T. orientalis maximised N resorption as N limitation increased (increasing NRE from 50.8 ± 3.3% to 71.7 ± 7.4%; reducing NRP from 0.70 ± 0.12% to 0.36 ± 0.13%) but did not alter PRE or PRP, whereas the S-strategist Schoenus pauciflorus maximised P resorption as P limitation increased (increasing PRE from 48.0 ± 5.6% to 73.5 ± 10.1%; reducing PRP from 0.053 ± 0.008% to 0.015 ± 0.004%) but did not alter NRE or NRP. These results show that the tissue N:P ratio and its associated δ¹⁵N enrichment are highly responsive indicators of the relative availability of N and P at the site and community level. However, they are not indicators of species-specific physiological requirements for N and P, or of likely responses of individual species to N or P enrichment, which are better interpreted from indicators such as RE and RP that describe nutrient retention behaviour.     

Nutritional value and intake of aquatic ferns (Azolla filiculoides Lam. and Salvinia molesta Mitchell.) in sows

Aquatic ferns (AFs) such as Azolla filiculoides and Salvinia molesta are grown on swine lagoons in the tropics and used in diets for pigs. The present work is aimed at evaluating their potential as feed ingredients for sows. When presented with ad libitum AFs, gilts weighing 110 ± 14 kg (mean ± SD), were able to ingest 9.1–9.7 kg fresh AF per day (from 597 to 630 g dry matter (DM) per day) and from 1240 to 1428 g DM per day when presented in a dry, ground form. A digestibility study was conducted, using sows weighing 213 ± 9 kg (mean ± SD), which were fed diets containing maize, soybean meal and 0, 150 or 300 g AF kg⁻¹ diet. The presence of AFs had a negative impact on the faecal digestibility of the crude protein, NDF and energy content of the whole diet (P<0.001) and on the ileal protein digestibility, especially with 300 g AFs kg⁻¹ diet. The level of AFs in the diet had no effect on stomach weight (P>0.05) but increased the weight of the rest of the gastrointestinal tract (P<0.001). The rate of AF fibre fermentation in the pig large intestine was measured using an in vitro gas test. The rates were much lower than tropical tree foliage, which can also be used in pig diets in the tropics. This could partly explain the low apparent digestibility of AFs in pigs. In conclusion, the inclusion level of AFs in rations for sows should be limited to 150 g AFs kg⁻¹ diet due to the low digestibility and energy density, as well as the negative impact on the digestibility of the whole diet.     

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.