Cooling streams with riparian trees: Thermal regime depends on total solar radiation penetrating the canopy

Riparian planting is often recommended for stream restoration, notably to mitigate solar heating of stream waters. However, plant leaves shade photosynthetically active radiation (PAR, 400–700 nm) more efficiently than the near-infrared (NIR, 700–3000 nm) wavelengths that comprise about half of incident solar radiation and so contribute equally to water heating. Total solar radiation (NIR + PAR) exposure at the reach scale is needed to avoid bias when predicting the thermal response of streams. In this article, we alert stream ecologists and riparian restorers to the water heating contribution of NIR and provide a means to account for both NIR and PAR in total solar radiation penetrating riparian canopies. We used spectral pyranometers to simultaneously measure total solar radiation and its NIR component under different tree canopies over a wide range of shade conditions as indexed by PAR exposure. Measurements were made during full overcast (diffuse lighting) conditions so as to ‘sample’ transmission of radiation through the complete canopy. The NIR proportion of sub-canopy solar radiation increased steadily with increasing shade and was appreciably greater under willow than pine canopies, while NZ native broadleaf rainforest had intermediate (rather variable) NIR content. Our trend lines for different tree canopies permit total sub-canopy solar radiation to be estimated from PAR exposure for unbiased modelling of stream thermal regimes.     

Experimental shading alters leaf litter breakdown in streams of contrasting riparian canopy cover

1. Headwater stream ecosystems are primarily heterotrophic, with allochthonous organic matter being the dominant energy. However, sunlight indirectly influences ecosystem structure and functioning, affecting microbial and invertebrate consumers and, ultimately, leaf litter breakdown. We tested the effects of artificial shading on litter breakdown rates in an open‐canopy stream (high ambient light) and a closed‐canopy stream (low ambient light). We further examined the responses of invertebrate shredders and aquatic hyphomycetes to shading to disentangle the underlying effects of light availability on litter breakdown. 2. Litter breakdown was substantially slower for both fast‐decomposing (alder, Alnus glutinosa) and slow‐decomposing (beech, Fagus sylvatica) leaf litters in artificially shaded stream reaches relative to control (no artificial shading) reaches, regardless of stream type (open or closed canopy). 3. Shredder densities were higher on A. glutinosa than on F. sylvatica litter, and shading had a greater effect on reducing shredder densities associated with A. glutinosa than those associated with F. sylvatica litter in both stream types. Fungal biomass was also negatively affected by shading. Results suggest that the effects of light availability on litter breakdown rates are mediated by resource quality and consumer density. 4. Results from feeding experiments, where A. glutinosa litter incubated under ambient light or artificial shade was offered to the shredder Gammarus fossarum, suggest that experimental shading and riparian canopy openness influenced litter palatability interactively. Rates of litter consumption by G. fossarum were decreased by experimental shading in the open‐canopy stream only. 5. The results suggest that even small variations in light availability in streams can mediate substantial within‐stream heterogeneity in litter breakdown. This study provides further evidence that changes in riparian vegetation, and thus light availability, influence organic matter processing in heterotrophic stream ecosystems through multiple trophic levels.     

Leaf structure of Syzygium spp. (Myrtaceae) in relation to site affinity within a tropical rain forest

This study examined four species of Syzygium (S. firmum, S. makul, S. operculatum, S. rubicundum) Myrtaceae, a tree genus that dominates the canopy of rain forests of south‐west Sri Lanka. Syzygium spp. occupy differing habitats with relation to succession and forest topography. We examined differences in leaf morphology and physiology in response to amount of shade, an important environmental variable affecting Syzygium distribution within the forest. To study change in leaf structure and physiology, environmental shelters were constructed simulating forest shade that differed in quality, quantity and duration. Seedlings were exposed to: (i) 0% shade (full sun, FS), red : far red (R : FR) ratio 1.27; (ii) 65% shade (large opening, LO) with direct sunlight similar to the centre of a large canopy opening, R : FR ratio 1.27; (iii) 82% shade (small opening, SO) with direct sunlight similar to the centre of a small canopy opening, R : FR ratio 1.27; (iv) 58% uniform light shade (LS) with a quality similar to the outside edge of a large canopy opening, R : FR ratio 1.05; (v) 85% uniform medium shade (MS) with a quality similar to the inside forest edge of a large canopy opening, R : FR ratio 0.97; (vi) 99% uniform deep shade (DS) similar to that of the forest understorey, R : FR ratio 0.23. The shelters were constructed in a large open area at the field station of the Sinharaja World Heritage site, Sri Lanka. Seedlings of each species were grown for two years in their respective shade treatments before physiological, morphological and anatomical measurements were made on leaves. Variation in leaf structure and physiology between the species was associated with differences in shade‐tolerance and water‐use. All species increased in photosynthesis rates and dimensions in leaf structure (leaf blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll) with decrease in shade. In contrast, stomatal conductivity was highest in the DS (99% shade) treatment. Leaves of Syzygium firmum were thickest and largest in area. S. firmum also had highest photosynthesis in the SO (82% shade) treatment. S. firmum was the most shade‐tolerant of all species: it grows well in low shade and its leaf structure suggests it to be the most conservative in water‐use of the Syzygium spp. In the forest S. firmum can persist in the forest shade as established seedlings, but grows best within canopy openings of late‐seral rain forest. Leaves of S. operculatum were thinnest but had highest stomatal densities of the four species. S. operculatum is considered shade‐intolerant, with a leaf structure suggesting it to be prone to desiccation, and by implication susceptible to drought. S. operculatum is found along streams within early seral rain forest habitat, often originating on stream banks after land clearance for cultivation. In the FS (0% shade) treatment, S. rubicundun had highest photosynthesis rates and greatest number of leaves but smallest leaf area of the Syzygium species. S. rubicundum is more shade‐intolerant but more efficient in water‐use than S. operculatum. S. rubicundum is a mid‐seral canopy tree of the midslope stands that are thought to have originated after catastrophic windthrows or swidden cultivation. The leaf physiology and structure of S. makul suggests it to be both moderately shade‐tolerant and conservative in water‐use. It is the most widely distributed Syzygium species across the topography of late‐seral rain forest. We suggest forest disturbance and hydrology are important environmental factors that influence distribution of Syzygium species across the topography. Results from this study contribute to a body of knowledge suggesting that canopy tree species of rain forests in south‐west Sri Lanka have discrete affinities to topography and differences in successional status, and that adaptations in leaf structure and physiology are indicative of such phenomena. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141 , 365–377.     

Causal Connections between Water Quality and Land Use in a Rural Tropical Island Watershed

We examined associations between riparian canopy cover, presence or absence of cattle, rainfall, solar radiation, month of year, dissolved oxygen, turbidity, salinity, and Enterococcus concentrations in riparian surface soils with Enterococcus geometric mean in-stream water concentrations at Waipā watershed on the north side of the Hawaiian island Kaua’i. Each 1% decrease in riparian canopy cover was associated with a 4.6 most probable number (MPN)/100 ml increase of the geometric mean of Enterococcus in stream water (P < 0.05). Each unit decrease in salinity (ppt) was associated with an increase of Enterococcus by 68.2 MPN/100 ml in-stream water geometric mean concentrations (P < 0.05). Month of year was also associated with increases in stream water Enterococcus geometric mean concentrations (P < 0.05). Reducing riparian canopy cover is associated with Enterococcus increases in stream water, suggesting that decreasing riparian vegetation density could increase fecal bacteria surface runoff.     

Effects of Stream Restoration on Woody Riparian Vegetation of Southern Appalachian Mountain Streams,North Carolina,U.S.A

Riparian revegetation, such as planting woody seedlings or live stakes, is a nearly ubiquitous component of stream restoration projects in the United States. Though evaluations of restoration success usually focus on in‐stream ecosystems, in order to understand the full impacts of restoration the effects on riparian ecosystems themselves must be considered. We examined the effects of stream restoration revegetation measures on riparian ecosystems of headwater mountain streams in forested watersheds by comparing riparian vegetation structure and composition at reference, restored, and degraded sites on nine streams. According to mixed model analysis of variance (ANOVA), there was a significant effect of site treatment on riparian species richness, basal area, and canopy cover, but no effect on stem density. Vegetation characteristics at restored sites differed from those of reference sites according to all metrics (i.e. basal area, canopy cover, and species composition) except species richness and stem density. Restored and degraded sites were structurally similar, with some overlap in species composition. Restored sites were dominated by Salix sericea and Cornus amomum (species commonly planted for revegetation) and a suite of disturbance‐adapted species also dominant at degraded sites. Differences between reference and restored sites might be due to the young age of restored sites (average 4 years since restoration), to reassembly of degraded site species composition at restored sites, or to the creation of a novel anthropogenic ecosystem on these headwater streams. Additional research is needed to determine if this anthropogenic riparian community type persists as a resilient novel ecosystem and provides valued riparian functions.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

Nitrate depletion in the riparian zone and stream channel of a small headwater catchment

A mass balance procedure was used to determine rates of nitrate depletion in the riparian zone and stream channel of a small New Zealand headwater stream. In all 12 surveys the majority of nitrate loss (56–100%) occurred in riparian organic soils, despite these soils occupying only 12% of the stream's border. This disproportionate role of the organic soils in depleting nitrate was due to two factors. Firstly, they were located at the base of hollows and consequently a disproportionately high percentage (37–81%) of the groundwater flowed through them in its passage to the stream. Secondly, they were anoxic and high in both denitrifying enzyme concentration and available carbon. Direct estimates ofin situ denitrification rate for organic soils near the upslope edge (338 mg N m^–2 h^–1) were much higher than average values estimated for the organic soils as a whole (0.3–2.1 mg N m^–2 h^–1) and suggested that areas of these soils were limited in their denitrification activity by the supply of nitrate. The capacity of these soils to regulate nitrate flux was therefore under-utilized. The majority of stream channel nitrate depletion was apparently due to plant uptake, with estimates of thein situ denitrification rate of stream sediments being less than 15% of the stream channel nitrate depletion rate estimated by mass balance.This study has shown that catchment hydrology can interact in a variety of ways with the biological processes responsible for nitrate depletion in riparian and stream ecosystems thereby having a strong influence on nitrate flux. This reinforces the view that those seeking to understand the functioning of these ecosystems need to consider hydrological phenomena.     

Predictions of stream nutrient and sediment yield changes following restoration of forested riparian buffers

Riparian tree planting is widely recognised as a means to improve water quality and stream habitat. However, shading of riparian pasture grasses can lead to channel widening, and riparian shade may limit the growth of macrophytes and algae that assimilate dissolved nutrients from the water column. We investigated concerns that riparian management could lead to increased yields of nutrients and sediments through a conceptual modelling exercise. A simple model of the trade-off between interception of nutrients in runoff by forest buffers versus reduction of in-stream uptake due to shade, predicted that a buffer strip alongside a small headwater stream would reduce nutrient export, while a buffer strip instigated as an isolated patch alongside a larger stream (c. >2.5 km² upstream catchment size) would increase nutrient export, as the relative amount of nutrients trapped by the buffer decreases as the nutrient load present in the stream water increases. However, in these larger streams with width exceeding approximately 6 m, sufficient light may reach the streambed for plant and algal growth, which in turn would promote instream nutrient processing. At the peak of streambank erosion after planting, predicted total sediment yield (hillslope plus bank sources) was appreciably higher than the hillslope pasture yield, but sediment yield stabilised c. 35–40 years after planting. When planting was extended over 40 years in the model, the sediment yield never exceeded that in pasture before planting. This conceptual modelling exercise shows that riparian tree planting programmes should commence in the headwaters and progress downstream to avoid nutrient yield increases. Significant sediment yield from bank stored sediment of small streams can be expected until the channel reaches the more stable, original forested width, but progressive planting may decrease the peak loads of sediment.     

Light environment under Rhododendron maximum thickets and estimated carbon gain of regenerating forest tree seedlings

Canopy tree recruitment is inhibited by evergreen shrubs in many forests. In the southern Appalachian mountains of the USA, thickets of Rhododendron maximum L. restrict dominant canopy tree seedling survival and persistence. Using R. maximum as a model system, we examined available light under the thickets and the photosynthetic responses of seedlings of canopy tree species. We tested the hypothesis that the additional shading from under R. maximum drives carbon gain in seedlings below the threshold for growth and survival. A reduction in light under the thicket was found where canopy openness (derived from canopy photographs) under R. maximum was half the amount measured in forest without R. maximum. R.␣maximum also reduced direct radiation by 50% and diffuse radiation by 12–29% compared to forest without the shrub layer. Mean mid-day PPFD (photosynthetically active photon flux density between 1000 and 1400 h) under R. maximum (obtained from quantum sensors) was below 10 mol m⁻² s⁻¹ on both clear and overcast days and the amount of sunflecks greater than 10 mol m⁻² s⁻¹ PPFD was only 0–20 min per day. In contrast, forest without R. maximum received a mean PPFD of 18–25 mol m⁻² s⁻¹ on clear days and a cumulative sunfleck duration of 100–220 min per day in all sky conditions. Consistent with light availability between the sites, daily carbon gain in Quercus rubra L. seedlings was lower in forest with R. maximum compared to forest where the shrub was absent. The presence of the shrub layer also significantly suppressed average mid-day photosynthesis of both Q. rubra and Prunus serotina Ehrt. seedlings on 8 out of 11 measurement dates. However, parameters derived from light response curves between seedlings growing in forest sites with or without a thicket of R. maximum was significantly different only in A _max (maximum photosynthetic rate), indicating a lack of further acclimation to the deeper shade under R. maximum. While the additional shade cast by R. maximum is sufficient to prevent the regeneration of tree seedlings under this shrub, there was sufficient heterogeneity in light under the thicket to imply that deep shade only partially explains the complete inhibition of regenerating canopy trees under R. maximum.     

Characterization of Shade Avoidance Responses in Lotus japonicus

Sessile plants must continuously adjust their growth and development to optimize photosynthetic activity under ever-fluctuating light conditions. Among such light responses in plants, one of the best-characterized events is the so-called shade avoidance, for which a low ratio of the red (R):far-red (FR) light intensities is the most prominent stimulus. Such shade avoidance responses enable plants to overtop their neighbors, thereby enhancing fitness and competitiveness in their natural habitat. Considerable progress has been achieved during the last decade in understanding the molecular mechanisms underlying the shade avoidance responses in the model rosette plant, Arabidopsis thaliana. We characterize here the fundamental aspects of the shade avoidance responses in the model legume, Lotus japonicus, based on the fact that its phyllotaxis (or morphological architecture) is quite different from that of A. thaliana. It was found that L. japonicus displays the characteristic shade avoidance syndrome (SAS) under defined laboratory conditions (a low R:FR ratio, low light intensity, and low blue light intensity) that mimic the natural canopy. In particular, the outgrowth of axillary buds (i.e., both aerial and cotyledonary shoot branching) was severely inhibited in L. japonicus grown in the shade. These results are discussed with special emphasis on the unique aspects of SAS observed with this legume.     

Drainage Size, Stream Intermittency, and Ecosystem Function in a Sonoran Desert Landscape

Understanding the interactions between terrestrial and aquatic ecosystems remains an important research focus in ecology. In arid landscapes, catchments are drained by a channel continuum that represents a potentially important driver of ecological pattern and process in the surrounding terrestrial environment. To better understand the role of drainage networks in arid landscapes, we determined how stream size influences the structure and productivity of riparian vegetation, and the accumulation of organic matter (OM) in soils beneath plants in an upper Sonoran Desert basin. Canopy volume of velvet mesquite (Prosopis velutina), as well as overall plant cover, increased along lateral upland–riparian gradients, and among riparian zones adjacent to increasingly larger streams. Foliar δ¹³C signatures for P. velutina suggested that landscape patterns in vegetation structure reflect increases in water availability along this arid stream continuum. Leaf litter and annual grass biomass production both increased with canopy volume, and total aboveground litter production ranged from 137 g m⁻² y⁻¹ in upland habitat to 446 g m⁻² y⁻¹ in the riparian zone of the perennial stream. OM accumulation in soils beneath P. velutina increased with canopy volume across a broad range of drainage sizes; however, in the riparian zone of larger streams, flooding further modified patterns of OM storage. Drainage networks represent important determinants of vegetation structure and function in upper Sonoran Desert basins, and the extent to which streams act as sources of plant-available water and/or agents of fluvial disturbance has implications for material storage in arid soils.     

The environmental preferences of three species of Australian freshwater fish in relation to the effects of riparian degradation

To assess the likely effects on three coexisting species of Australian freshwater fish of riparian loss, we examined the temperature, light, and habitat preferences of each species in relation to commonly documented effects of riparian degradation on stream environments. Such effects include reduced shade, instream structure, and water depth and increased temperature and invasive instream vegetation. Xiphophorus helleri, Gambusia holbrooki, and Melanotaenia duboulayi differed significantly in mean swimming depth, preferences for light and habitat, and in their patterns of behavioural change through the day. Values of interspecific spatial overlap (25–58%) indicated that the overall environmental preferences of G. holbrooki, X. helleri, and M. duboulayi were reasonably distinct. Habitat alterations associated with riparian removal are likely to favour the exotic species G. holbrooki over the native species M. duboulayi, but the results for X. helleri suggest that not all poeciliid␣species are strong indicators of degraded conditions.     

Effects of riparian canopy cover on salmonid diet and prey selectivity in low nutrient streams

This study focuses on stream sections within a relatively low nutrient catchment in south‐east Ireland in an attempt to characterize the probable effects of riparian canopy on salmonid diet and prey selectivity within two size classes of nursery stream. Sampling found that brown trout Salmo trutta diet changed significantly in response to riparian canopy regardless of stream size. The observation that S. trutta within unshaded stream sites did not feed on drifting terrestrial prey items to the same extent as those within shaded streams was not due to a lack of availability of this food source. There was no evidence to suggest that S. trutta selectively choose particular prey items.     

Use of digital webcam images to track spring green-up in a deciduous broadleaf forest

Understanding relationships between canopy structure and the seasonal dynamics of photosynthetic uptake of CO₂ by forest canopies requires improved knowledge of canopy phenology at eddy covariance flux tower sites. We investigated whether digital webcam images could be used to monitor the trajectory of spring green-up in a deciduous northern hardwood forest. A standard, commercially available webcam was mounted at the top of the eddy covariance tower at the Bartlett AmeriFlux site. Images were collected each day around midday. Red, green, and blue color channel brightness data for a 640 × 100-pixel region-of-interest were extracted from each image. We evaluated the green-up signal extracted from webcam images against changes in the fraction of incident photosynthetically active radiation that is absorbed by the canopy (f _APAR), a broadband normalized difference vegetation index (NDVI), and the light-saturated rate of canopy photosynthesis (A _max), inferred from eddy flux measurements. The relative brightness of the green channel (green %) was relatively stable through the winter months. A steady rising trend in green % began around day 120 and continued through day 160, at which point a stable plateau was reached. The relative brightness of the blue channel (blue %) also responded to spring green-up, although there was more day-to-day variation in the signal because blue % was more sensitive to changes in the quality (spectral distribution) of incident radiation. Seasonal changes in blue % were most similar to those in f _APAR and broadband NDVI, whereas changes in green % proceeded more slowly, and were drawn out over a longer period of time. Changes in A _max lagged green-up by at least a week. We conclude that webcams offer an inexpensive means by which phenological changes in the canopy state can be quantified. A network of cameras could offer a novel opportunity to implement a regional or national phenology monitoring program.     

基于人体健康的3种城市森林夏季林内紫外线辐射环境特征比较研究

以中波紫外线为主的紫外辐射对人体健康具有多种生物学影响,城市森林能够为居民提供温和的紫外辐射环境。为了解林下紫外线辐射环境特征是否存在树种间差异,对北京市3种常见遮荫树种的夏季林下紫外辐射(UV)强度、林内与林外UV辐射的比值(SR)、UV-B在总UV辐射中占比(UV-B/UV),以及VD合成和红斑效应两种人体作用有效辐射强度(UV_VD、UV_er)进行了测算。结果表明：(1)三种林分林内紫外辐射总量是林外的3%—10%,不同林分的林内UV强度具有显著差异,元宝枫林对UV屏蔽能力最强,其次是栾树林和国槐林;(2)三种林分林冠对不同波长上紫外辐射能量的屏蔽能力具有明显的一致性,林冠对UV-B的屏蔽能力没有在UVA波段强和稳定,林内UV-B/UV普遍高于林外,其中元宝枫林最高,其次为栾树林,国槐林最低;(3)林冠明显改变了日光UV_VD和UV_er两种人体作用光谱曲线的形态,三种林分内的人体作用光谱曲线形态相似,强度上,林内外UV_VD/UV_er值均接近1,不同林分间...     

The expansion of woody riparian vegetation,and subsequent stream restoration,influences the metabolism of prairie streams

1. Tallgrass prairies and their streams are highly endangered ecosystems, and many remaining streams are threatened by the encroachment of woody riparian vegetation. An increase in riparian vegetation converts the naturally open‐canopy prairie streams to closed‐canopy systems. The effects of a change in canopy cover on stream metabolism are unknown. 2. Our goal was to determine the effects of canopy cover on prairie stream metabolism during a 4‐year period in Kings Creek, KS, U.S.A. Metabolic rates from forested reaches were compared to rates in naturally open‐canopy reaches and restoration reaches, the latter having closed canopies in 2006 and 2007 and open canopies in 2008 and 2009. Whole‐stream metabolism was estimated using the two‐station diurnal method. Chlorophyll a concentrations and mass of filamentous algae were measured after riparian removal to assess potential differences in algal biomass between reaches with open or closed canopies. 3. Metabolic rates were spatially and temporally variable even though the sites were on very similar streams or adjacent to each other within streams. Before riparian vegetation removal, whole‐stream community respiration (CR) and net ecosystem production were greater with greater canopy cover. In the vegetation removal reaches, gross primary production was slightly greater after removal. 4. Chlorophyll a concentrations were marginally significantly greater in open (naturally open and removal reaches) than in closed canopy and differed significantly between seasons. Filamentous algal biomass was greater in open than in closed‐canopy reaches. 5. Overall, the restoration allowed recovery of some features of open‐canopy prairie streams. Woody expansion apparently increases CR and moves prairie stream metabolism towards a more net heterotrophic state. An increase in canopy cover decreases benthic chlorophyll, decreases dominance of filamentous algae and potentially alters resources available to the stream food web. The results of this study provide insights for land managers and conservationists interested in preserving prairie streams in their native open‐canopy state.     

Theories on adaptations to high light intensity in the aquatic moss Fontinalis

Abstract

Fontinalis antipyretica, collected from a cold stream in bright sunlight, served as a model for adaptations of Fontinalis to sun and shade. Sun leaves produced red pigments and had less chlorophyll, greater specific leaf weight, and less specific leaf area than shade leaves. The chlorophyll a/b ratio was not significantly different under high and low light conditions. Except for the relatively constant a/b ratio, Fontinalis behaved like higher plants in its sun/shade differences.     

Impaired phytochrome-mediated shade-avoidance responses in the aurea mutant of tomato

Internode extension-growth responses to neighbouring plants and to red to far-red ratios (R:FR) were investigated in wild-type (WT) and aurea (au)-mutant seedlings of tomato grown under natural radiation. The genomic location of the au mutant is not known, but one of its consequences is the reduced phytochrome level. In WT seedlings, internode growth was promoted by the presence of non-shading neighbours reflecting far-red light (FR), the shade of a tall canopy, FR provided as a supplement during the photoperiod, and FR pulses either provided at the end of the day or delayed into the dark period. Supplementary FR during the photoperiod also promoted growth in herbicide-treated partially bleached WT seedlings. The au mutant showed higher background extension-growth rates, but only responded to the most severe treatments: deep shade light and very low R:FR at the end of the day, i.e. au-mutant seedlings were less sensitive than WT seedlings to R:FR signals. Wild-type seedlings were transferred from the glasshouse to a growth room and exposed to white light with two levels of phytochrome-absorbable radiation but similar phytochrome photoequilibria and radiation for photosynthesis. The plants exposed to the lowest level showed a transient increase of internode extension growth rate and a simultaneous reduction of response to FR pulses, i.e. reproduced some of the features of au-mutant seedlings. Phytochrome itself could set the degree of response to Pfr during neighbour detection.     

Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes

Summary Plant canopy shade reduces photosynthetic photon flux density (PPFD) and ratio of red to far-red light (z). Both effects can cause plants to increase potential for light acquisition through vertical growth and leaf area expansion. Clonal plants such as Eichhornia crassipes might alternatively increase light interception via horizontal growth of stolons or rhizomes and placement of new ramets in less shaded microsites. Effect of simulated canopy shade and component effects of PPFD and z were tested by filtering or adding light uniformly, to a whole group of connected ramets, or locally, to individual ramets within a group. In uniform treatments, low PPFD reduced total growth but low z did not. Low PPFD and low z independently reduced stolon and ramet production and caused etiolation of petioles; effect of low PPFD plus low z on ramet production was greater than that of either factor alone. Lateral clonal growth thus did not seem to be a response to uniform shading; instead, uniformly low PPFD or low z increased partitioning to established ramets. Low z changed partitioning without changing total growth. In local treatments, reduction of growth of individual ramets due to low PPFD and inhibition of new ramet production attributable to spectral composition of light were mitigated when connected ramets were unshaded; plants may respond differently to patchy than to uniform shade.     

Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.