Environmental determinants of vegetation in the drawdown zones of a Columbia River Treaty reservoir: a template for ecosystem enhancement.

Water storage reservoirs alternately inundate and expose the drawdown zones, limiting riparian vegetation that provides wildlife habitats and contributes to the aquatic food-web. To characterize plant distributions and hydrogeomorphic associations, we inventoried quadrats in transects extending from the full-pool (FP) margin, downwards 12 m through the drawdown zones at sites around the Duncan Reservoir in British Columbia, Canada. Among the 69 plant species, black cottonwoods (Populus trichocarpa), willows (primarily Salix sitchensis) and other trees and shrubs occurred sparsely, rarely extending below 2 m below FP. Perennial herbaceous plants, especially horsetail (Equisetum arvense) and sedges (primarily Carex utriculata), were most common, extending down ~5 m below FP, and ruderal annual plants occurred sparsely at greater depths. Vegetation Cover and Species Richness were correlated with environmental factors, with (1) Elevation being highly influential, reflecting inundation duration and depth. (2) Position, longitudinal location, reflected greater vegetation diversity downstream of the reservoir. (3) Finer Substrate texture was favorable to retain moisture, but coarse sediments would resist erosion. (4) Shallow Slope was favorable to reduce drainage and included finer sediments. (5) Distance from the FP shoreline could reflect seed source proximity. Stepwise linear modeling with combined environmental factors accounted for ~30% of the variation in Vegetation Cover and Richness, and Canonical Correspondence Analysis revealed plant groupings relative to the environmental influences. At this and other storage reservoirs, regimes that reduce the frequency and duration of inundation could promote vegetation in locations with suitable environmental conditions in the upper drawdown zones, thus providing ecosystem enhancement.

How trees thrive in a dry climate: diurnal and seasonal hydrology and water relations in a riparian cottonwood grove.

In semi-arid ecoregions, trees are restricted to river valley floodplains where river water supplements the limited precipitation. To characterize the associated diurnal and seasonal dynamics in hydrology and water relations, we studied narrowleaf cottonwoods (Populus angustifolia) along a prairie river in Canada. From June through August, the shallow soil moisture was depleted but moisture remained higher above the alluvial groundwater table, which dropped to 2.3 m along with river recession. Throughout the summer, with the daily rise in temperature and insolation, foliar stomatal conductance (gs) and transpiration (E) increased to midday and then fell, thus maintaining the midday leaf water potential (Ψmd) above ~-1.7 MPa. This Ψmd approximated the water potential associated with 12% loss of xylem conductivity due to cavitation for branches (P12); the Ψmd and P12 varied independently across eight trees, providing differences in relative hydraulic risk. Sap flux density (Fd) was measured with thermal dissipation probes, and revealed diurnal patterns similar to foliar E. In contrast to our expectation, the daily Fd maxima were consistent through the summer despite the seasonal recession in water supply. Canopy conductances (GS), derived from Fd, sapwood area and canopy area, declined with vapor pressure deficit (D) and fell slightly in late summer along with stomatal sensitivity, which reflects the magnitude of decrease in GS with increasing D. For spatial up-scaling, satellite-derived near-infrared reflectance of vegetation revealed the woodland phenology, with leaf expansion from May through June and gradual decline in photosynthetic productivity through the summer. Thus, the phreatophytic cottonwoods: (i) sustained substantial water use and productivity through the warm and dry summer, by (ii) accessing shallow soil moisture and then deeper alluvial groundwater, and (iii) providing diurnal stomatal regulation, to (iv) avoid xylem cavitation and (v) maintain fairly constant hydraulic conductance.

Multiple processes contribute to methane emission in a riparian cottonwood forest ecosystem.

Methane emission from trees may partially or completely offset the methane sink in upland soils, the only process that has been regularly included in methane budgets for forest ecosystems. Our objective was to analyze multiple biogeochemical processes that influence the production, oxidation and transport of methane in a riparian cottonwood ecosystem and its adjacent river. We combined chamber flux measurements on tree stems, forest soil and the river surface with eddy covariance measurements of methane net ecosystem exchange. In addition, we tested whether methanogens were present in cottonwood stems, shallow soil layers and alluvial groundwater. Average midday peak in net methane emission measured by eddy covariance was c. 12 nmol m-2  s-1 . The average uptake of methane by soils (0.87 nmol m-2  s-1 ) was largely offset by tree stem methane emission (0.75 nmol m-2  s-1 ). There was evidence of methanogens in tree stems but not in shallow soil. Growing season (May-September) cumulative net methane emission (17.4 mmol CH4  m-2 ) included methane produced in cottonwood stems and methane input to the nocturnal boundary layer from the forest and the adjacent river. The multiple processes contributing to methane emission illustrated the linked nature of these adjacent terrestrial and aquatic ecosystems.

Prospective impacts of oil spills on floodplain vegetation: Both crude oil and diluted bitumen increase foliar temperatures, senescence and abscission in three cottonwood (Populus) species.

Oil pipelines are vulnerable at river crossings since floods can expose and rupture pipes, releasing oil that floats and coats floodplain vegetation. This study investigated the consequences of oil coatings on leaves of cottonwoods (riparian poplars), the predominant trees in floodplain woodlands around the Northern Hemisphere. The study compared conventional crude oil (CO) versus diluted bitumen (dilbit, DB), heavy oil originating from the Alberta oil sands; with petroleum jelly (PJ) as a reference. The treatments increased leaf surface temperatures (Tleaf) in narrowleaf and plains cottonwoods (Populus angustifolia, P. deltoides) and balsam poplars (P. balsamifera) (Control = 21.8°C, PJ = 23.7°C; CO = 26.2°C; DB = 28.1°C; Tair = 25°C). The leaf warming followed stomatal occlusion from the foliar coating, which would reduce transpiration and evaporative cooling, combined with increased solar warming with the darker oils. Tleaf varied across the three cottonwood species, with cooler, narrow, narrowleaf cottonwood leaves; intermediate plains cottonwood leaves; and warmer, darker, balsam poplar leaves (average Tleaf: narrowleaf = 23.8°C, plains = 24.3°C, and balsam = 26.7°C), with similar warming in each species following the different treatments. Across species and treatments, Tleaf was tightly correlated with foliar condition, which assessed turgor versus wilting of leaf blades and petioles, along with leaf necrosis and senescence (r2 = 0.980, narrowleaf; 0.998, plains; 0.852, balsam). This tight association indicates validity of both Tleaf and foliar condition as diagnostic measures. Crude oil and dilbit had similar foliar impacts, and for both, leaf abscission occurred within 2 to 3 weeks. Consequently, following an oil spill, remediation should commence quickly but extending vegetation removal beyond a few weeks would have limited benefit since the contaminated leaves would have abscised.

Gibberellins and Heterosis in Crops and Trees: An Integrative Review and Preliminary Study with Brassica.

Heterosis, or hybrid vigor, has contributed substantially to genetic improvements in crops and trees and its physiological basis involves multiple processes. Four associations with the phytohormone gibberellin (GA) indicate its involvement in the regulation of heterosis for shoot growth in maize, sorghum, wheat, rice, tomato and poplar. (1) Inbreds somewhat resemble GA-deficient dwarfs and are often highly responsive to exogenous GA3. (2) Levels of endogenous GAs, including the bioeffector GA1, its precursors GA19 and GA20, and/or its metabolite GA8, are higher in some fast-growing hybrids than parental genotypes. (3) Oxidative metabolism of applied [3H]GAs is more rapid in vigorous hybrids than inbreds, and (4) heterotic hybrids have displayed increased expression of GA biosynthetic genes including GA 20-oxidase and GA 3-oxidase. We further investigated Brassica rapa, an oilseed rape, by comparing two inbreds (AO533 and AO539) and their F1 hybrid. Seedling emergence was faster in the hybrid and potence ratios indicated dominance for increased leaf number, area and mass, and stem mass. Overdominance (heterosis) was displayed for root mass, leading to slight heterosis for total plant mass. Stem contents of GA19,20,1 were similar across the Brassica genotypes and increased prior to bolting; elongation was correlated with endogenous GA but heterosis for shoot growth was modest. The collective studies support a physiological role for GAs in the regulation of heterosis for shoot growth in crops and trees, and the Brassica study encourages further investigation of heterosis for root growth.

A Lightweight Leddar Optical Fusion Scanning System (FSS) for Canopy Foliage Monitoring.

A growing need for sampling environmental spaces in high detail is driving the rapid development of non-destructive three-dimensional (3D) sensing technologies. LiDAR sensors, capable of precise 3D measurement at various scales from indoor to landscape, still lack affordable and portable products for broad-scale and multi-temporal monitoring. This study aims to configure a compact and low-cost 3D fusion scanning system (FSS) with a multi-segment Leddar (light emitting diode detection and ranging, LeddarTech), a monocular camera, and rotational robotics to recover hemispherical, colored point clouds. This includes an entire framework of calibration and fusion algorithms utilizing Leddar depth measurements and image parallax information. The FSS was applied to scan a cottonwood (Populus spp.) stand repeatedly during autumnal leaf drop. Results show that the calibration error based on bundle adjustment is between 1 and 3 pixels. The FSS scans exhibit a similar canopy volume profile to the benchmarking terrestrial laser scans, with an r2 between 0.5 and 0.7 in varying stages of leaf cover. The 3D point distribution information from FSS also provides a valuable correction factor for the leaf area index (LAI) estimation. The consistency of corrected LAI measurement demonstrates the practical value of deploying FSS for canopy foliage monitoring.

A prescription for drug-free rivers: uptake of pharmaceuticals by a widespread streamside willow.

Following human excretion and limited removal with wastewater treatment, pharmaceuticals are accumulating in rivers worldwide. These chemicals can challenge the health of fish and aquatic organisms and since rivers provide drinking water sources, there is concern for cumulative exposure to humans. In this study, we discovered that sandbar willow (Salix exigua), a predominant riparian shrub along streams throughout North America, has the capacity to quickly remove pharmaceuticals from aqueous solutions. Our study tracked [3H]- or [14C]-labeled substances including 17α-ethynylestradiol (EE2), a synthetic estrogen in oral contraceptives; the antihypertensive, diltiazem (DTZ); and the anti-anxiety drug, diazepam (DZP); and for comparison, atrazine (ATZ), a root-absorbed herbicide. In growth chambers, willow saplings removed 40-80% of the substances from solutions in 24 h. Following uptake, the EE2 and DTZ were retained within the roots, while DZP and ATZ were partly passed on to the shoots. The absorbed EE2 was unextractable and apparently bound to the root tissue, while DTZ, DZP, and ATZ remained largely soluble (extractable). The uptake and translocation of the pharmaceuticals, reflected in the transpiration stream and root concentration factors, were reasonably predicted from their physicochemical properties, including octanol-water partitioning coefficients. These findings suggest the removal of pharmaceuticals as an unrecognized ecosystem service provided by riparian vegetation and especially the inundation tolerant sandbar willow. This encourages the conservation of riparian willows that line riverbanks, to remove pharmaceuticals and other contaminants. This phytoremediation also encourages the preservation of complex, braided channels and islands, which increase the extent of stream shorelines and riparian willows.

Heterosis in poplar involves phenotypic stability: cottonwood hybrids outperform their parental species at suboptimal temperatures.

Heterosis or hybrid vigor is common in hybrid poplars, and to investigate its occurrence and physiological basis we compared narrowleaf cottonwoods, Populus angustifolia James, prairie cottonwoods, Populus deltoides Bartr. ex Marsh, and their native intersectional hybrids, P. × acuminata Rydb., from Alberta, Canada. Clonal replicates from 10 separate trees from each taxon were raised in growth chambers at different temperatures (T). Growth was similarly vigorous across the taxa at 20 and 24 °C, and morphological and physiological traits of the hybrids were generally intermediate between the parental species, or similar to the larger parent, demonstrating additive inheritance or dominance, respectively. Growth declined at 18 and 15 °C particularly in the parental species, and consequently hybrid vigor was displayed for root and especially leaf growth. Stomatal distributions and chlorophyll indices were intermediate in the hybrids and unaffected by T. Foliar nitrogen (N), net assimilation (Asat), stomatal conductance (gs) and transpiration (E) per unit of leaf area were lower in the hybrids, but the hybrids generally had larger leaf areas. Water-use efficiencies (Asat/gs) were similar across the taxa and reduced with warming, while nitrogen-use efficiencies (Asat/N) increased. δ13C was correlated with leaf mass per area, which varied across the taxa. Photosynthesis (Asat) was correlated with chlorophyll content index, N and/or gs in P. deltoides and the hybrids, but not in P. angustifolia, indicating different physiological limitations. We conclude that heterosis in P. × acuminata results from the compound benefits from multiple dominant traits, and superior growth particularly at suboptimal conditions. This indicates phenotypic stability or environmental adaptability, whereby heterozygosity provides metabolic diversity that allows hybrids to thrive across a broader environmental range.

The Irrigation Effect: How River Regulation Can Promote Some Riparian Vegetation.

River regulation impacts riparian ecosystems by altering the hydrogeomorphic conditions that support streamside vegetation. Obligate riparian plants are often negatively impacted since they are ecological specialists with particular instream flow requirements. Conversely, facultative riparian plants are generalists and may be less vulnerable to river regulation, and could benefit from augmented flows that reduce drought stress during hot and dry periods. To consider this 'irrigation effect' we studied the facultative shrub, netleaf hackberry (Celtis reticulata), the predominant riparian plant along the Hells Canyon corridor of the Snake River, Idaho, USA, where dams produce hydropeaking, diurnal flow variation. Inventories of 235 cross-sectional transects revealed that hackberry was uncommon upstream from the reservoirs, sparse along the reservoir with seasonal draw-down and common along two reservoirs with stabilized water levels. Along the Snake River downstream, hackberry occurred in fairly continuous, dense bands along the high water line. In contrast, hackberry was sparsely scattered along the free-flowing Salmon River, where sandbar willow (Salix exigua), an obligate riparian shrub, was abundant. Below the confluence of the Snake and Salmon rivers, the abundance and distribution of hackberry were intermediate between the two upstream reaches. Thus, river regulation apparently benefited hackberry along the Snake River through Hells Canyon, probably due to diurnal pulsing that wets the riparian margin. We predict similar benefits for some other facultative riparian plants along other regulated rivers with hydropeaking during warm and dry intervals. To analyze the ecological impacts of hydropeaking we recommend assessing daily maxima, as well as daily mean river flows.

Bud phenology and growth are subject to divergent selection across a latitudinal gradient in Populus angustifolia and impact adaptation across the distributional range and associated arthropods.

Temperate forest tree species that span large geographical areas and climatic gradients often have high levels of genetic variation. Such species are ideal for testing how neutral demographic factors and climate-driven selection structure genetic variation within species, and how this genetic variation can affect ecological communities. Here, we quantified genetic variation in vegetative phenology and growth traits in narrowleaf cottonwood, Populus angustifolia, using three common gardens planted with genotypes originating from source populations spanning the species' range along the Rocky Mountains of North America (ca. 1700 km). We present three main findings. First, we found strong evidence of divergent selection (Q ST > F ST) on fall phenology (bud set) with adaptive consequences for frost avoidance. We also found evidence for selection on bud flush duration, tree height, and basal diameter, resulting in population differentiation. Second, we found strong associations with climate variables that were strongly correlated with latitude of origin. More strongly differentiated traits also showed stronger climate correlations, which emphasizes the role that climate has played in divergent selection throughout the range. We found population × garden interaction effects; for some traits, this accounted for more of the variance than either factor alone. Tree height was influenced by the difference in climate of the source and garden locations and declined with increasing transfer distance. Third, growth traits were correlated with dependent arthropod community diversity metrics. Synthesis. Overall, we conclude that climate has influenced genetic variation and structure in phenology and growth traits and leads to local adaptation in P. angustifolia, which can then impact dependent arthropod species. Importantly, relocation of genotypes far northward or southward often resulted in poor growth, likely due to a phenological mismatch with photoperiod, the proximate cue for fall growth cessation. Genotypes moved too far southward suffer from early growth cessation, whereas those moved too far northward are prone to fall frost and winter dieback. In the face of current and forecasted climate change, habitat restoration, forestry, and tree breeding efforts should utilize these findings to better match latitudinal and climatic source environments with management locations for optimal future outcomes.

Gravel-bed river floodplains are the ecological nexus of glaciated mountain landscapes.

Gravel-bed river floodplains in mountain landscapes disproportionately concentrate diverse habitats, nutrient cycling, productivity of biota, and species interactions. Although stream ecologists know that river channel and floodplain habitats used by aquatic organisms are maintained by hydrologic regimes that mobilize gravel-bed sediments, terrestrial ecologists have largely been unaware of the importance of floodplain structures and processes to the life requirements of a wide variety of species. We provide insight into gravel-bed rivers as the ecological nexus of glaciated mountain landscapes. We show why gravel-bed river floodplains are the primary arena where interactions take place among aquatic, avian, and terrestrial species from microbes to grizzly bears and provide essential connectivity as corridors for movement for both aquatic and terrestrial species. Paradoxically, gravel-bed river floodplains are also disproportionately unprotected where human developments are concentrated. Structural modifications to floodplains such as roads, railways, and housing and hydrologic-altering hydroelectric or water storage dams have severe impacts to floodplain habitat diversity and productivity, restrict local and regional connectivity, and reduce the resilience of both aquatic and terrestrial species, including adaptation to climate change. To be effective, conservation efforts in glaciated mountain landscapes intended to benefit the widest variety of organisms need a paradigm shift that has gravel-bed rivers and their floodplains as the central focus and that prioritizes the maintenance or restoration of the intact structure and processes of these critically important systems throughout their length and breadth.

Higher photosynthetic capacity from higher latitude: foliar characteristics and gas exchange of southern, central and northern populations of Populus angustifolia.

Narrowleaf cottonwood (Populus angustifolia James) is an obligate riparian poplar that is a foundation species in river valleys along the Rocky Mountains, spanning 16° of latitude from southern Arizona, USA to southern Alberta, Canada. Its current distribution is fragmented, and genetic variation shows regional population structure consistent with the effects of geographic barriers and past climate. It is thus very well-suited for investigating ecophysiological adaptation associated with latitude. In other section Tacamahaca poplar species, genotypes from higher latitudes show evidence of short-season adaptation with foliar traits that contribute to higher photosynthetic capacity. We tested for similar adaptation in three populations of narrowleaf cottonwoods: from Arizona (south), Alberta (north) and Utah, near the centre of the latitudinal distribution. We propagated 20 genotypes from each population in a common garden in Alberta, and measured foliar and physiological traits after 3 years. Leaves of genotypes from the northern population had higher leaf mass per area (LMA), increased nitrogen (N) content and higher carotenoid and chlorophyll content, and these were associated with higher light-saturated net photosynthesis (Asat). In leaves of all populations the majority of stomata were abaxial, with the proportion of abaxial stomata highest in the southern population. Stomatal conductance (gs) and transpiration rates were higher in the northern population but water-use efficiency (Asat/gs) and leaf carbon isotope composition (δ(13)C) did not differ across the populations. These results (i) establish links between Asat and gs, N, chlorophyll and LMA among populations within this species, (ii) are consistent with the discrimination of populations from prior investigation of genetic variation and (iii) support the concept of latitudinal adaptation, whereby deciduous trees from higher latitudes display higher photosynthetic capacity, possibly compensating for a shorter and cooler growth season and reduced insolation.

Biological effects and toxicity of diluted bitumen and its constituents in freshwater systems.

Approximately 50 billion cubic meters of bitumen resides within the oil sands region of Alberta, Canada. To facilitate the transport of bitumen from where it is extracted to where it is processed, the bitumen is diluted with natural gas condensate ('dilbit'), synthetic crude from hydrocracking bitumen ('synbit'), or a mixture of both ('dilsynbit'). A primary consideration for the effects of diluted bitumen products on freshwater organisms and ecosystems is whether it will float on the water surface or sink and interact with the stream or lake sediments. Evidence from a spill near Kalamazoo, MI, in 2010 and laboratory testing demonstrate that the nature of the spill and weathering of the dilbit, synbit or dilsynbit prior to and during contact with water will dictate whether the product floats or sinks. Subsequent toxicological data on the effects of dilbit and other diluted bitumen products on freshwater organisms and ecosystems are scarce. However, the current literature indicates that dilbit or bitumen can have significant effects on a wide variety of toxicological endpoints. This review synthesizes the currently available literature concerning the fate and effects of dilbit and synbit spilled into freshwater, and the effects of bitumen and bitumen products on aquatic organisms and ecosystems. Dilbit is likely to provide ecological impacts that are similar to and extend from those that follow from exposure to lighter crude oil, but the prospect of bitumen settling after binding to suspended sediments elevates the risk for benthic impacts in streams and lakes.

Floodplain forest succession reveals fluvial processes: A hydrogeomorphic model for temperate riparian woodlands.

River valley floodplains are physically-dynamic environments where fluvial processes determine habitat gradients for riparian vegetation. These zones support trees and shrubs whose life stages are adapted to specific habitat types and consequently forest composition and successional stage reflect the underlying hydrogeomorphic processes and history. In this study we investigated woodland vegetation composition, successional stage and habitat properties, and compared these with physically-based indicators of hydraulic processes. We thus sought to develop a hydrogeomorphic model to evaluate riparian woodland condition based on the spatial mosaic of successional phases of the floodplain forest. The study investigated free-flowing and dam-impacted reaches of the Kootenai and Flathead Rivers, in Idaho and Montana, USA and British Columbia, Canada. The analyses revealed strong correspondence between vegetation assessments and metrics of fluvial processes indicating morphodynamics (erosion and shear stress), inundation and depth to groundwater. The results indicated that common successional stages generally occupied similar hydraulic environments along the different river segments. Comparison of the spatial patterns between the free-flowing and regulated reaches revealed greater deviation from the natural condition for the braided channel segment than for the meandering segment. This demonstrates the utility of the hydrogeomorphic approach and suggests that riparian woodlands along braided channels could have lower resilience than those along meandering channels and might be more vulnerable to influences such as from river damming or climate change.

Development of a spatially-distributed hydroecological model to simulate cottonwood seedling recruitment along rivers.

Dam operations have altered flood and flow patterns and prevented successful cottonwood seedling recruitment along many rivers. To guide reservoir flow releases to meet cottonwood recruitment needs, we developed a spatially-distributed, GIS-based model that analyzes the hydrophysical requirements for cottonwood recruitment. These requirements are indicated by five physical parameters: (1) annual peak flow timing relative to the interval of seed dispersal, (2) shear stress, which characterizes disturbance, (3) local stage recession after seedling recruitment, (4) recruitment elevation above base flow stage, and (5) duration of winter flooding, which may contribute to seedling mortality. The model categorizes the potential for cottonwood recruitment in four classes and attributes a suitability value at each individual spatial location. The model accuracy was estimated with an error matrix analysis by comparing simulated and field-observed recruitment success. The overall accuracies of this Spatially-Distributed Cottonwood Recruitment model were 47% for a braided reach and 68% for a meander reach along the Kootenai River in Idaho, USA. Model accuracies increased to 64% and 72%, respectively, when fewer favorability classes were considered. The model predicted areas of similarly favorable recruitment potential for 1997 and 2006, two recent years with successful cottonwood recruitment. This model should provide a useful tool to quantify impacts of human activities and climatic variability on cottonwood recruitment, and to prescribe instream flow regimes for the conservation and restoration of riparian woodlands.

Hydrologic linkages between a climate oscillation, river flows, growth, and wood Δ13C of male and female cottonwood trees.

To investigate climatic influence on floodplain trees, we analysed interannual correspondences between the Pacific Decadal Oscillation (PDO), river and groundwater hydrology, and growth and wood (13)C discrimination (Δ(13)C) of narrowleaf cottonwoods (Populus angustifolia) in a semi-arid prairie region. From the Rocky Mountain headwaters, river discharge (Q) was coordinated with the PDO (1910-2008: r(2) = 0.46); this pattern extended to the prairie and was amplified by water withdrawal for irrigation. Floodplain groundwater depth was correlated with river stage (r(2) = 0.96), and the cottonwood trunk basal area growth was coordinated with current- and prior-year Q (1992-2008: r(2) = 0.51), increasing in the mid-1990s, and decreasing in 2000 and 2001. Annual Δ(13)C decreased during low-flow years, especially in trees that were higher or further from the river, suggesting drought stress and stomatal closure, and male trees were more responsive than females (-0.86 versus -0.43‰). With subsequently increased flows, Δ(13)C increased and growth recovered. This demonstrated the linkages between hydroclimatic variation and cottonwood ecophysiology, and we conclude that cottonwoods will be vulnerable to drought from declining river flows due to water withdrawal and climate change. Trees further from the river could be especially affected, leading to narrowing of floodplain forests along some rivers.

Green revolution trees: semidwarfism transgenes modify gibberellins, promote root growth, enhance morphological diversity, and reduce competitiveness in hybrid poplar.

Semidwarfism has been used extensively in row crops and horticulture to promote yield, reduce lodging, and improve harvest index, and it might have similar benefits for trees for short-rotation forestry or energy plantations, reclamation, phytoremediation, or other applications. We studied the effects of the dominant semidwarfism transgenes GA Insensitive (GAI) and Repressor of GAI-Like, which affect gibberellin (GA) action, and the GA catabolic gene, GA 2-oxidase, in nursery beds and in 2-year-old high-density stands of hybrid poplar (Populus tremula × Populus alba). Twenty-nine traits were analyzed, including measures of growth, morphology, and physiology. Endogenous GA levels were modified in most transgenic events; GA(20) and GA(8), in particular, had strong inverse associations with tree height. Nearly all measured traits varied significantly among genotypes, and several traits interacted with planting density, including aboveground biomass, root-shoot ratio, root fraction, branch angle, and crown depth. Semidwarfism promoted biomass allocation to roots over shoots and substantially increased rooting efficiency with most genes tested. The increased root proportion and increased leaf chlorophyll levels were associated with changes in leaf carbon isotope discrimination, indicating altered water use efficiency. Semidwarf trees had dramatically reduced growth when in direct competition with wild-type trees, supporting the hypothesis that semidwarfism genes could be effective tools to mitigate the spread of exotic, hybrid, and transgenic plants in wild and feral populations.

Sand and sandbar willow: a feedback loop amplifies environmental sensitivity at the riparian interface.

Riparian or streamside zones support dynamic ecosystems with three interacting components: flowing water, alluvia (river-transported sediments), and vegetation. River damming influences all three, and subsequent responses can provide insight into underlying processes. We investigated these components along the 315-km Hells Canyon corridor of the Snake River that included reaches upstream, along, and downstream from three large dams and reservoirs, and along the Salmon River, a free-flowing tributary. Sandbar willow was generally the woody plant at the lowest bank position and was abundant along upstream reaches (53, 45, 67% of transects), sparse along reservoirs (11, 12, 0%), and sparse along the Snake River downstream (11%). It was prolific along the undammed Salmon River (83%) and intermediate along the Snake River below the Salmon inflow (27%), indicating partial recovery with the contribution of water and sediments. Along these rivers, it commonly occurred on sandy substrates, especially on shallow-sloped surfaces, and emerged from interstitial sands between cobbles on steeper surfaces. However, along the Snake River below the dams, sandbars have eroded and willows were sparse on remnant, degrading sand surfaces. We conclude that a feedback loop exists between sands and sandbar willow. Sand favors willow colonization and clonal expansion, and reciprocally the extensively branched willows create slack-water zones that protect and trap sands. This feedback may sustain surface sands and sandbar willows along free-flowing river systems and it amplifies their mutual vulnerability to river damming. Following damming, sediment-depleted water is released downstream, eroding surface sands and reducing willow colonization and expansion. With willow decline, sands are further exposed and eroded, compounding these impacts. From this feedback, we predict the coordinated depletion of surface sands and riparian willows along dammed rivers throughout the Northern Hemisphere.

Streamside trees: responses of male, female and hybrid cottonwoods to flooding.

Cottonwoods, riparian poplars, are dioecious and prior studies have indicated that female poplars and willows can be more abundant than males in low-elevation zones, which are occasionally flooded. We investigated the response to flooding of clonal saplings of 12 male and 9 female narrowleaf cottonwoods (Populus angustifolia) grown for 15 weeks in a greenhouse, along with three females of a co-occurring native hybrid (Populus × jackii = Populus deltoides × Populus balsamifera). Three water-level treatments were provided, with substrate inundation as the flood treatment. In the non-flooded condition, the hybrids produced about four-fold more dry weight (DW) than the narrowleaf cottonwoods (P < 0.01). In both cottonwood taxa, flooding reduced stem height and DW, root and leaf area and weight, leaf chlorophyll and stomatal conductance (all P < 0.01). Inundation increased the foliar carbon-to-nitrogen ratio (+11%; P < 0.05) but did not significantly alter leaf water potential (mean -1.5 MPa), or foliar δ(13)C, which was lower in P. angustifolia (-32.8‰) than P. × jackii (-31.5‰; P < 0.05). Water level influenced the root distribution as roots were sparse in the saturated substrate and abundant in the capillary fringe above. The male and female P. angustifolia genotypes grew similarly with the favorable water levels, but the males tended to be more inhibited by flooding. Sapling DW of males was reduced by 56% compared with a 44% reduction for females (P = 0.1), and there were similar lower reductions for leaf, stem and root DW in females. These results demonstrate the inundation response of floodplain trees and suggest relative flood tolerance as: P. angustifolia female > P. angustifolia male > P. × jackii female. This indicates that narrowleaf cottonwoods are relatively flood tolerant and suggests that females are more flood tolerant than males. We propose the concept of 'strategic positioning', whereby the seed-producing females could be better adapted to naturally flooded, low-elevation streamside zones where seedling recruitment generally occurs.

Favorable fragmentation: river reservoirs can impede downstream expansion of riparian weeds.

River valleys represent biologically rich corridors characterized by natural disturbances that create moist and barren sites suitable for colonization by native riparian plants, and also by weeds. Dams and reservoirs interrupt the longitudinal corridors and we hypothesized that this could restrict downstream weed expansion. To consider this "reservoir impediment" hypothesis we assessed the occurrences and abundances of weeds along a 315-km river valley corridor that commenced with an unimpounded reach of the Snake River and extended through Brownlee, Oxbow, and Hells Canyon reservoirs and dams, and downstream along the Snake River. Sampling along 206 belt transects with 3610 quadrats revealed 16 noxious and four invasive weed species. Ten weeds were upland plants, with Canada thistle (Cirsium arvense) restricted to the upstream reaches, where field morning glory (Convolvulus arvensis) was also more common. In contrast, St. John's wort (Hypericum perforatum) was more abundant below the dams, and medusahead wildrye (Taeniatherum caput-medusae) occurred primarily along the reservoirs. All seven riparian species were abundant in the upstream zones but sparse or absent below the dams. This pattern was observed for the facultative riparian species, poison hemlock (Conium maculatum) and perennial pepperweed (Lepidium latifolium), the obligate riparian, yellow nut sedge (Cyperus esculentus), the invasive perennial, reed canary grass (Phalaris arundinacea), and three invasive riparian trees, Russian olive (Elaeagnus angustifolia), false indigo (Amorpha fruticosa), and tamarisk (Tamarix spp.). The hydrophyte purple loosestrife (Lythrum salicaria) was also restricted to the upstream zone. These longitudinal patterns indicate that the reservoirs have impeded the downstream expansion of riparian weeds, and this may especially result from the repetitive draw-down and refilling of Brownlee Reservoir that imposes a lethal combination of drought and flood stress. The dams and reservoirs may also interrupt hydrochory, the downstream flow of seeds and clonal fragments. We thus conclude that with some operational patterns, dams and reservoirs can impede the downstream expansion of riparian weeds.