Burial depth distribution of fennel pondweed tubers (Potamogeton pectinatus) in relation to foraging by Bewick's swans

Deep burial in the sediment of tubers of fennel pondweed (Potamogeton pectinatus) has been explained in terms of avoidance by escape against consumption by Bewick's swans (Cygnus columbianus bewickii) in autumn. We therefore expected changes in foraging pressure to ultimately result in a change in the tuber distribution across sediment depth. A trade-off underlies this idea: deep tubers are less accessible to swans but must be larger to meet the higher energy demands of sprouting in spring. To test this prediction, we compared tuber burial depth over a gradient of foraging pressure both across space and across time. Tuber samples were obtained after aboveground plant senescence but before arrival of Bewick's swans. First, we compared the current tuber bank depth profile in a shallow lake with high foraging pressure, the Lauwersmeer, with that in two wetlands with moderate and low foraging pressure. Second, we compared the current tuber burial in the Lauwersmeer with that in the early 1980s when exploitation by swans had just started there. In accordance with our hypothesis, we found significantly deeper burial of tubers under high consumption risk compared to low consumption risk, both when comparing sites and comparing time periods. Since tubers in effect only survive to the next spring, the observed differences in burial depth among sites and over time cannot be a direct result of tuber losses due to consumption by swans. Rather, these observations suggest adaptive responses in tuber burial related to foraging pressure from Bewick's swans in the recent past. We thus propose that fennel pondweed exhibits flexible avoidance by escape, of a kind rarely described for plants, where both phenotypic plasticity and genotype sorting may contribute to the observed differences in tuber burial.     

Reduced tuber banks of fennel pondweed due to summer grazing by waterfowl

In this study we investigated the effect of summer bird herbivory on the belowground tuber formation of fennel pondweed (Potamogeton pectinatus L.). Cumulative grazing pressure of four waterfowl species (mute swans, mallards, gadwalls and coots) in the summer was calculated based on timing of grazing and body mass of the grazers. The resulting grazing pressures were significantly negatively correlated with mean autumn tuber biomass in three of the four years of study. Moreover, summer grazing pressures explained more of the variance in tuber densities than water depth, sediment particle size distribution or any interactions of these variables did in the same three years. We propose that herbivory early in the summer has the most substantial impact on the clonal reproduction of macrophytes. Herbivores with a large body mass and early congregation for moulting may be the key waterfowl species in diminishing propagule biomass. Hence, they may present pre-emptive, time-staggered competition to consumers of the belowground biomass in autumn, such as migratory swans and diving ducks.     

The effect of herbivores on genotypic diversity in a clonal aquatic plant

In clonal plants, vegetative parts may outcompete seeds in the absence of disturbance, limiting the build‐up of genotypic diversity through repeated seedling recruitment (RSR). Herbivory may provide disturbance and trigger establishment of strong colonizers (seeds) at the expense of strong competitors (clonal propagules). In the clonal aquatic fennel pondweed Potamogeton pectinatus, two distinct herbivore guilds may modify the dynamics of propagation. In winter, Bewick's swans may deplete patches of tubers, promoting seedling establishment in spring. In summer, seed consumption by waterfowl can reduce the density of viable seeds but grazing may also reduce tuber production and hence facilitate seedling establishment. This study is among the first to experimentally test herbivore impact on plant genotypic diversity. We assess the separate and combined effects of both herbivore guilds on genotypic diversity and structure of fennel pondweed beds. Using microsatellites, we genotyped P. pectinatus from an exclosure experiment and assessed the contribution of herbivory, dispersal and sexual reproduction to the population genetic structure. Despite the predominance of clonal propagation in P. pectinatus, we found considerable genotypic diversity. Within the experimental blocks, kinship among genets decreased with geographic distance, clearly identifying a role for RSR in the maintenance of genotypic diversity within the fennel pondweed beds. However, over a period of five years, none of the herbivory treatments affected genotypic diversity. Hence, sexual reproduction on a local scale is important in this putatively clonal plant and possibly sufficient to ensure a relatively high genotypic diversity even in the absence of herbivores. Although we cannot preclude a role of herbivory in shaping genotypic diversity of a clonal plant, after five years of exclusion of the two investigated herbivore guilds no measurable effect on genotypic diversity was detected.     

Overcompensation and grazing optimisation in a swan–pondweed system?

1. The general notion is that negative effects of vertebrate herbivores on water plants, which play a key role in freshwaters, prevail, and that positive feedbacks of herbivores on plants are insignificant. 2. The most likely systems to find such positive feedbacks are those in which herbivores exert strong feeding pressures on plants during part of the year. Previous theoretical work has suggested that compensatory production occurs when migratory Bewick's swans forage on tubers of fennel pondweed. As a corollary, the swans can exploit the tubers down to a level that maximises their tuber yield. 3. In order to test these hypotheses, I measured pondweed tuber biomass on three occasions per year (just before and after foraging, and just before tuber sprouting) in three consecutive years. The 17 sampling sites in the Lauwersmeer (the Netherlands) were classified according to their silt content and water depth. Within four silt‐depth classes, I predicted for each year tuber biomass production and, from that, the optimum foraging threshold that would result in the maximum tuber biomass yield. 4. Water depth did not affect tuber production, and silt content only did in one of the 3 years. In accordance with overcompensation predictions, tuber production was higher at plots with moderate foraging pressures than at plots with little or no grazing. However, the winter and summer conditions following the swan foraging had large unpredictable effects on tuber mortality and production. 5. These results indicate that overcompensation by fennel pondweed occurs and that Bewick's swans are generally able to profit from it, albeit without fine‐tuning of the foraging threshold to the yield.     

Above- and below-ground vertebrate herbivory may each favour a different subordinate species in an aquatic plant community

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species.     

Swan foraging shapes spatial distribution of two submerged plants,favouring the preferred prey species

Compared to terrestrial environments, grazing intensity on belowground plant parts may be particularly strong in aquatic environments, which may have great effects on plant-community structure. We observed that the submerged macrophyte, Potamogeton pectinatus, which mainly reproduces with tubers, often grows at intermediate water depth and that P. perfoliatus, which mainly reproduces with rhizomes and turions, grows in either shallow or deep water. One mechanism behind this distributional pattern may be that swans prefer to feed on P. pectinatus tubers at intermediate water depths. We hypothesised that when swans feed on tubers in the sediment, P. perfoliatus rhizomes and turions may be damaged by the uprooting, whereas the small round tubers of P. pectinatus that escaped herbivory may be more tolerant to this bioturbation. In spring 2000, we transplanted P. perfoliatus rhizomes into a P. pectinatus stand and followed growth in plots protected and unprotected, respectively, from bird foraging. Although swan foraging reduced tuber biomass in unprotected plots, leading to lower P. pectinatus density in spring 2001, this species grew well both in protected and unprotected plots later that summer. In contrast, swan grazing had a dramatic negative effect on P. perfoliatus that persisted throughout the summer of 2001, with close to no plants in the unprotected plots and high densities in the protected plots. Our results demonstrate that herbivorous waterbirds may play a crucial role in the distribution and prevalence of specific plant species. Furthermore, since their grazing benefitted their preferred food source, the interaction between swans and P. pectinatus may be classified as ecologically mutualistic.     

Differences in tolerance of pondweeds and charophytes to vertebrate herbivores in a shallow Baltic estuary

It has been suggested that herbivorous waterfowl may be important in shaping aquatic plant communities in shallow wetlands. As such, a shift from canopy forming pondweeds to bottom-dwelling charophytes in a formerly turbid pondweed dominated lake has been partly attributed to waterfowl herbivory. Here we study the separate and combined effects of both belowground herbivory in spring by whooper swans and Bewick ‘s swans, and grazing in summer by waterfowl and fish on the community composition in a shallow Baltic estuary during one year. The macrophyte community was dominated by charophytes (mainly Chara aspera) with Potamogeton pectinatus and Najas marina present as subdominants. Other species were rare. Both spring and summer herbivory had no effect on total plant biomass. However, P. pectinatus was more abundant in plots that were closed to spring and summer herbivores. N. marina was more abundant in grazed plots, whereas Chara spp. biomass remained unaffected. Probably belowground propagules of both C. aspera and P. pectinatus were consumed by swans but since C. aspera bulbils were numerous it may have compensated for the losses. P. pectinatus may not have fully recovered from foraging on tubers and aboveground biomass. Our results are in line with other studies in Chara dominated lakes, which found no effect of grazing on summer aboveground Chara biomass, whereas several studies report strong effects of herbivory in lakes dominated by P. pectinatus.     

Foraging costs and accessibility as determinants of giving-up densities in a swan-pondweed system

We measured the patch use behaviour of Bewick's swans (Cygnus columbianus bewickii) feeding on below ground tubers of fennel pondweed (Potamogeton pectinatus). We compared the swans’ attack rates, foraging costs and giving‐up densities (GUDs) in natural and experimental food patches that differed in water depth. Unlike most studies that attribute habitat‐specific differences in GUDs to predation risk, food quality or foraging substrate, we quantified the relative importance of energetic costs and accessibility. Accessibility is defined as the extent to which the animal's morphology restricts its harvest of all food items within a food patch. Patch use behaviours were measured at shallow (ca 0.4 m) and deep (ca 0.6 m) water depths on sandy sediments. In a laboratory foraging experiment, when harvesting food patches, the swan's attack rate (m³ s^?1) did not differ between depths. In deep water the energetic costs of surfacing, feeding and trampling were 1.13 to 1.21 times higher than in shallow water with a tendency to spend relatively more time trampling, the most expensive activity. Taking time allocation as measured in the field into account, foraging in deep water was 1.26 times as expensive as in shallow water. In the lake the GUD in shallow water was on average 12.9 g m^?2. If differences in energetic costs were the only factor determining differences in GUDs, then the deep water GUD should be 14.2 g m^?2. Instead, the mean GUD in deep water was 20.2 g m^?2, and therefore energetic costs explain just 18% of the difference in GUDs. At deep sites, 24% of tuber biomass was estimated to be out of reach, and we calculated a maximum accessible foraging depth of 0.86 m. This is close to the published 0.84 m based on body measurements. A laboratory experiment with food offered at a depth of 0.89 m confirmed that it was just out of reach. The agreement between calculated and observed maximum accessible foraging depths suggests that accessibility largely explains the remaining difference in GUDs with depth, and it confirms the existence of partial prey refuges in this system.     

Influence of photoperiod on growth,pigment composition and vegetative propagule formation for Potamogeton nodosus Poir. and Potamogeton pectinatus L.

Photoperiods of 10 or 12 h enhanced vegetative propagule production by Potamogeton nodosus Poir. and P. pectinatus L. The proportion of plants producing vegetative propagules and the number and weight of propagules per plant increased over time, and were greatest after 8 weeks. Maximum vegetative propagule production was 12±6 tubers per plant (mean±S.D.; n=7) for P. pectinatus and 5±3 for P. nodosus. Tuber or winter-bud dry weight accounted for 38 or 27% of total plant dry weight, respectively, for plants grown for 56 days under a 10-h photoperiod. Vegetative propagules appeared to be produced instead of new shoot tissue under short-day conditions. Chlorophyll a and carotenoid content were lower for P. nodosus at short photoperiods, but not for P. pectinatus. The changes in chlorophyll a:b and carotenoid:ratio chlorophyll a suggest that synthesis/degradation of these pigments is tightly coupled and that short photoperiods may initiate senescence in P. nodosus.     

Control of potato tuber dormancy and sprouting by expression of sense and antisense genes of pyrophosphatase in potato

Inorganic pyrophosphate (PPi) is an enzyme involved in sugar metabolism in potato tubers. In our previous study, we isolated an inorganic pyrophosphatase (PPase) gene from potato and obtained the transgenic potato plants transformed with the sense and antisense PPase genes respectively. In the present experiment, the physiological indexes, tuber dormancy, and sprouting characteristics of the transgenic potatoes were analyzed and evaluated. The result showed that the PPase activity and the inorganic phosphate content of tubers were lower in the antisense transgenic plant lines but were higher in the sense transgenic plant lines, compared with wild-type tubers. Soluble sugars, such as glucose, fructose and sucrose increased in transgenic plants that had overexpression of the sense PPase gene, but decreased in the antisense transgenic plant lines, compared with wild-type tubers. Tuber sprouting time of the antisense transgenic plants were delayed for 2 and 3 weeks and reached the 100 % sprouting rate only after 14 and 16 weeks storage compared with the wild-type when tubers are stored under 25 and 4 °C, respectively. In contrast, tuber sprouting time of the sense transgenic plants was earlier by approximately 2 weeks than that of wild-type tubers under these storage temperatures.     

The use of a flexible patch leaving rule under exploitative competition: a field test with swans

Learning animals are predicted to use a flexible patch-leaving threshold (PLT) while foraging in a depletable environment under exploitative competition. This prediction was tested in flock-feeding Bewick's swans ( Cygnus columbianus bewickii ) depleting hidden tubers of fennel pondweed ( Potamogeton pectinatus ) in a two-dimensional, continuous environment. The swans' patch residence time was measured by combining recordings of the foraging behaviour and movement paths. The tuber biomass density was measured before and after the period of exploitation, using the presumable foraging window of the swans as the scale of measurement. Swan foraging was simulated in order to predict the effects of flexible and fixed PLTs, respectively, on the patch residence time and the spatial heterogeneity of the tuber biomass density. Flexible PLTs were predicted to lead to short and decreasing patch residence times and a decrease in the coefficient of variation in tuber biomass densities, whereas the reverse was generally the case for fixed PLTs. Observed patch residence times did not decrease with time and were intermediate between those predicted for swans with flexible and fixed PLTs. Furthermore, an increase of the coefficient of variation in the tuber biomass density was observed. Given the observed giving-up biomass densities the most likely model was one with swans with a fixed rather than a flexible PLT. These results point at factors that may affect the spacing behaviour or constrain the use of a flexible PLT in swans.     

Contribution of insectivorous avifauna to top down control of Lindera benzoin herbivores at forest edge and interior habitats

Predation of herbivorous Lepidoptera larvae by insectivorous avifauna was estimated on Lindera benzoin in edge and interior habitats at two sites in eastern Pennsylvania (USA). Clay baits modeled after Epimecis hortaria (Geometridae) larvae, the primary herbivore of L. benzoin at our study sites, were used to estimate predation by birds. In both habitat types, models were placed on uninjured L. benzoin leaves as well as on leaves that had prior insect herbivore damage. Rates of model attack were greater, and model longevity reduced, in forest edge plots compared to interiors. Naturally occurring herbivore damage on L. benzoin was greater in forest interiors. However, model attack was not significantly greater on leaves with prior herbivory damage, suggesting that birds do not effectively use this type of leaf damage as a cue in their foraging. Our findings are consistent with a contribution of bird predation towards top-down control of herbivory in this system. We further discuss these results in a broader context considering the possible effects of habitat type on leaf quality, leaf defense, and herbivore performance.     

Propagule banks, herbivory and nutrient supply control population development and dominance patterns in macroalgal blooms

Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis . Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella . Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella . Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha , thereby negatively affecting Pilayella . Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.     

Influence of migrant tundra swans (Cygnus columbianus) and Canada geese (Branta canadensis) on aquatic vegetation at Long Point, Lake Erie, Ontario

Numerous studies have shown that large, herbivorous waterfowl can reduce quantity of aquatic plants during the breeding or wintering season, but relatively few document herbivory effects at staging areas. This study was done to determine if feeding activities of tundra swans (Cygnus columbianus columbianus) and Canada geese (Branta canadensis) had a measurable additive influence on the amount of aquatic plants, primarily muskgrass (Chara vulgaris), wild celery (Vallisneria americana), and sago pondweed (Potamogeton pectinatus), removed during the fall migration period at Long Point, Lake Erie, Ontario. Exclosure experiments done in fall 1998 and 1999 showed that, as compared to ducks and abiotic factors, these two large herbivorous waterfowl did not have any additional impact on above or below ground biomass of those aquatic plants. As expected, however, there were substantial seasonal reductions in above-ground and below-ground biomass of aquatic plants in wetlands that were heavily used by all waterfowl. We suggest that differences in large- and small-scale habitat use, feeding activity, and food preferences between tundra swans and other smaller waterfowl as well as compensatory herbivory contributed to our main finding that large waterfowl did not increase fall reductions of Chara spp, V. Americana, and P. pectinatus biomass.     

Nature of enhanced respiration during sprouting of aged potato seed-tubers

Respiration of 18-month-old Solarium tuberosum L. tubers was about 53% greater than that of 6-month-old tubers during sprouting at 23°C; yet, a significant loss of sprout vigor in the older tubers was apparent. Involvement of alternative oxidase (AO) in the age-induced difference in tuber respiration was assessed. AO was only detected in immunoblots if tissue disks from tubers were pre-incubated for 24 h prior to isolation of submitochondrial membrane particles (SMPs). No AO¹ was detected in SMPs from nonincubated tuber tissue of either age, indicating that it was not contributing to tuber respiration during sprouting as previously thought. Respiratory control and ADP/O ratios indicated that oxidative phosphorylation was fully coupled to electron transport in mitochondria isolated from 6- and 18-month-old tubers. Cytochrome c oxidase (EC 1.9.3.1) activities of intact mitochondria were also not affected by tuber age. The difference in respiration during sprouting was unique to whole tubers, as oxygen consumption by mitochondria from young and oid tubers was equal on a milligram protein basis. Sprouting 18-month-old tubers had 15% more mitochondrial protein per gram fresh weight than did 6-month-old tubers. Older tubers also produced more ATP than younger tubers prior to and during sprouting, through a fully coupled, Cyt-mediated respiratory pathway, reduced sprout vigor notwithstanding. From 5 to 10 days of sprouting, coinciding with development of the age-induced difference in whole-tuber respiration, ATP concentration in 18-month-old tubers increased to become 52% higher than that in 6-month-old tubers. ATP synthase (EC 3.6.1.34), assessed by SDS-PAGE and immunoblots of β- and oligomycin-sensitivity conferring protein-subunits, also increased as a proportion of SMP protein in older tubers during this period. Relative to 6-month-old tubers, the increased respiration and associated oxidative phosphorylation of 18-rnonth-old tubers during sprouting were probably in response to a lower adenylate energy charge (AEC) prior to sprouting (from 0 fo 5 days). From 5 to 10 days of sprouting, AEC of 18-rnonth-old tubers increased to equal that of 6-month-old tubers and the two tuber ages maintained the same AEC for the remainder of the 20-day sprouting interval. Higher respiration and lower AEC of older tubers in storage at 4°C, along with the fact that older tubers respired at a higher rate to achieve the same AEC as younger tubers during sprouting, indicate greater utilization of ATP by older tubers.     

Effects of storage conditions on sprouting of microtubers of yam (Dioscorea cayenensis–D. rotundata complex)

The control of field tuber dormancy in the yam (Dioscorea cayenensis–D. rotundata complex) is poorly understood. Although studies have examined single environmental factors and chemical treatments that might prolong tuber dormancy and storage, only a few were focused on further tuber sprouting. The present study concerns microtubers obtained by in vitro culture. When microtubers were harvested (after 9 months of culture) and directly transferred on a new medium without hormones, the tubers rapidly sprouted in in vitro conditions. No dormancy was observed in this case. Harvested microtubers were also stored dry in jars in sterile conditions during 2 to 18 weeks before in vitro sprouting. In this case, microtubers stored during 18 weeks sprouted more rapidly than those stored 8 weeks. A constant “dormancy-like period” (storage duration + sprouting delay) was observed, between 20 and 28 weeks respectively for the more rapid and the slower microtubers. The size of the tubers used for the storage had great influence on further sprouting. The larger they were, the better they sprouted. Light during storage had no effect on the sprouting delay while a temperature of 25 °C permit a quicker sprouting than 18 °C. The medium used to obtain microtubers could also have an effect on sprouting rate.     

Seasonally changing habitat use patterns among roving herbivorous fishes in the southern Red Sea: the role of temperature and algal community structure

Coral reefs are characterized by intense herbivory. Spatial patterns in herbivory—particularly along the depth gradient—influence the distribution and abundance of algae. Depth gradients in herbivorous reef fishes are generally assumed to be temporally stable, but this assumption has rarely been questioned. Here, we use underwater visual census and herbivore exclusion experiments to study the community composition and temporal patterns in habitat use by roving herbivorous fishes in an environment characterized by profound seasonal changes in algal biomass and distribution and extreme summer temperatures. Among the 18 species of roving herbivores recorded, parrotfishes were dominant in species richness and biomass, while regional endemic species represented 77 % of the total biomass. During most of the year, roving herbivores aggregate in the shallow reef zones and their biomass declines with depth. The herbivore community on the reef flat is distinct from that in deeper zones. The former is characterized by Siganus rivulatus, Acanthurus gahhm and Hipposcarus harid, while the deeper reef zones are characterized by S. ferrugineus, Chlorurus sordidus and Ctenochaetus striatus. In summer, the distinct community structures among reef zones are lost as reef flat herbivores tend to exploit deeper reef zones and some reef crest species venture on to the reef flat. This summer change in herbivore distribution is also reflected in reduced turf biomass and increased yield to herbivores in the deeper reef zones. Habitat use is related to the feeding mode such that browsers dominate the reef flat and scrapers the reef crest, while the seasonal changes correspond to changes in availability of targeted algal resources. These seasonal changes appear to be driven by the extreme temperatures in summer, reaching 36 °C on the shallow reef flat.     

Regulation of potato tuber dormancy and sprouting

Dormancy is the final stage of tuber life serving to preserve tubers as organs of vegetative reproduction under unfavorable growth conditions. Since the duration of potato tuber dormancy and their sprouting time have significant economic importance, much attention is given to the study of the regulation of these processes. This review considers metabolite, genetic, and hormonal aspects of regulation of potato (Solanum tuberosum L.) tuber dormancy and sprouting. Particular attention is paid to the relationship between processes occurring in different parts of the tuber: its storage tissues and buds. The interaction of hormonal and metabolite (carbohydrate) regulation of dormancy and sprouting is discussed.     

Separate and interactive effects of competition and herbivory on the growth, expansion, and tuber formation of Hydrilla verticillata

This study examined the interaction and main-effect impacts of herbivory by the leaf-mining fly Hydrellia pakistanae and plant competition from Vallisneria americana on the growth, expansion and tuber formation of Hydrilla verticillata in a 2 × 2 factorial design experiment. The study was conducted in 14,000-L tanks, over two growing seasons. Each tank represented a single experimental unit and contained 32 1-L pots. At the beginning of the experiment half of these were planted with H. verticillata while the other half were either left empty or planted with V. americana (the competitor). H. pakistanae fly larvae (the herbivore) were added to tanks as appropriate. No significant interactions were identified between herbivory and competition on any parameter of H. verticillata growth analyzed (i.e., total tank biomass accumulation, total number of rooting stems, total tuber number, total tuber mass, and tuber size), indicating that the factors were operating independently and neither antagonism nor synergism was occurring. Both competition and herbivory impacted the growth of H. verticillata. H. verticillata plants grown in the presence of V. americana developed less total biomass, had fewer total basal stems, had fewer tubers and less tuber mass per tank, and produced significantly smaller tubers relative to control plants. Herbivory also suppressed H. verticillata biomass accumulation and tended to suppress the number and total mass of tubers produced in each tank. Both factors showed 30–40% reduction of total H. verticillata biomass, although the mechanism of impact was different. Competition suppressed expansion of H. verticillata into adjoining pots but had little impact on its growth in pots where it was originally planted. Herbivory resulted in a general suppression of growth of H. verticillata in all pots. Although herbivory significantly impacted H. verticillata biomass, it did not result in competitive release for V. americana under the current experimental conditions. We conclude that management activities that promote competition or herbivory will impact the growth and expansion of H. verticillata. Furthermore, since these factors operated independently, the combined use of both factors should be beneficial for suppression of H. verticillata dominance.     

The Influence of Growth Regulators on Sprouting of Hydrilla Tubers and Turions

The effects of different growth regulators on the sproutingof hydrilla reproductive propagules (Hydrilla verticillata (L.F.)Rovle) were studied. Some growth regulators either had no effector inhibited sprouting. Ethephon, gibberellic acid, and thioureaincreased tuber sprouting significantly. Thiourea was less effectivein inducing sprouting in turions than in tubers. Maintainingtubers at 5 ± 2°C enhanced sprouting significantly.Tubers harvested in summer responded differently to growth regulatorsthan some of the tubers harvested in winter. Therefore, it wasconcluded that two types of dormancy exist in hydrilla tubers.