Plant chlorophyll fluorescence: active and passive measurements at canopy and leaf scales with different nitrogen treatments.

Most studies assessing chlorophyll fluorescence (ChlF) have examined leaf responses to environmental stress conditions using active techniques. Alternatively, passive techniques are able to measure ChlF at both leaf and canopy scales. However, the measurement principles of both techniques are different, and only a few datasets concerning the relationships between them are reported in the literature. In this study, we investigated the potential for interchanging ChlF measurements using active techniques with passive measurements at different temporal and spatial scales. The ultimate objective was to determine the limits within which active and passive techniques are comparable. The results presented in this study showed that active and passive measurements were highly correlated over the growing season across nitrogen treatments at both canopy and leaf-average scale. At the single-leaf scale, the seasonal relation between techniques was weaker, but still significant. The variability within single-leaf measurements was largely related to leaf heterogeneity associated with variations in CO2 assimilation and stomatal conductance, and less so to variations in leaf chlorophyll content, leaf size or measurement inputs (e.g. light reflected and emitted by the leaf and illumination conditions and leaf spectrum). This uncertainty was exacerbated when single-leaf analysis was limited to a particular day rather than the entire season. We concluded that daily measurements of active and passive ChlF at the single-leaf scale are not comparable. However, canopy and leaf-average active measurements can be used to better understand the daily and seasonal behaviour of passive ChlF measurements. In turn, this can be used to better estimate plant photosynthetic capacity and therefore to provide improved information for crop management.

Sun-induced fluorescence - a new probe of photosynthesis: First maps from the imaging spectrometer HyPlant.

Variations in photosynthesis still cause substantial uncertainties in predicting photosynthetic CO2 uptake rates and monitoring plant stress. Changes in actual photosynthesis that are not related to greenness of vegetation are difficult to measure by reflectance based optical remote sensing techniques. Several activities are underway to evaluate the sun-induced fluorescence signal on the ground and on a coarse spatial scale using space-borne imaging spectrometers. Intermediate-scale observations using airborne-based imaging spectroscopy, which are critical to bridge the existing gap between small-scale field studies and global observations, are still insufficient. Here we present the first validated maps of sun-induced fluorescence in that critical, intermediate spatial resolution, employing the novel airborne imaging spectrometer HyPlant. HyPlant has an unprecedented spectral resolution, which allows for the first time quantifying sun-induced fluorescence fluxes in physical units according to the Fraunhofer Line Depth Principle that exploits solar and atmospheric absorption bands. Maps of sun-induced fluorescence show a large spatial variability between different vegetation types, which complement classical remote sensing approaches. Different crop types largely differ in emitting fluorescence that additionally changes within the seasonal cycle and thus may be related to the seasonal activation and deactivation of the photosynthetic machinery. We argue that sun-induced fluorescence emission is related to two processes: (i) the total absorbed radiation by photosynthetically active chlorophyll; and (ii) the functional status of actual photosynthesis and vegetation stress.

Vertical gradient in soil temperature stimulates development and increases biomass accumulation in barley.

We have detailed knowledge from controlled environment studies on the influence of root temperature on plant performance, growth and morphology. However, in all studies root temperature was kept spatially uniform, which motivated us to test whether a vertical gradient in soil temperature affected development and biomass production. Roots of barley seedlings were exposed to three uniform temperature treatments (10, 15 or 20°C) or to a vertical gradient (20-10°C from top to bottom). Substantial differences in plant performance, biomass production and root architecture occurred in the 30-day-old plants. Shoot and root biomass of plants exposed to vertical temperature gradient increased by 144 respectively, 297%, compared with plants grown at uniform root temperature of 20°C. Additionally the root system was concentrated in the upper 10cm of the soil substrate (98% of total root biomass) in contrast to plants grown at uniform soil temperature of 20°C (86% of total root biomass). N and C concentrations in plant roots grown in the gradient were significantly lower than under uniform growth conditions. These results are important for the transferability of 'normal' greenhouse experiments where generally soil temperature is not controlled or monitored and open a new path to better understand and experimentally assess root-shoot interactions.

Remote sensing of plant-water relations: An overview and future perspectives.

Vegetation is a highly dynamic component of the Earth surface and substantially alters the water cycle. Particularly the process of oxygenic plant photosynthesis determines vegetation connecting the water and carbon cycle and causing various interactions and feedbacks across Earth spheres. While vegetation impacts the water cycle, it reacts to changing water availability via functional, biochemical and structural responses. Unravelling the resulting complex feedbacks and interactions between the plant-water system and environmental change is essential for any modelling approaches and predictions, but still insufficiently understood due to currently missing observations. We hypothesize that an appropriate cross-scale monitoring of plant-water relations can be achieved by combined observational and modelling approaches. This paper reviews suitable remote sensing approaches to assess plant-water relations ranging from pure observational to combined observational-modelling approaches. We use a combined energy balance and radiative transfer model to assess the explanatory power of pure observational approaches focussing on plant parameters to estimate plant-water relations, followed by an outline for a more effective use of remote sensing by their integration into soil-plant-atmosphere continuum (SPAC) models. We apply a mechanistic model simulating water movement in the SPAC to reveal insight into the complexity of relations between soil, plant and atmospheric parameters, and thus plant-water relations. We conclude that future research should focus on strategies combining observations and mechanistic modelling to advance our knowledge on the interplay between the plant-water system and environmental change, e.g. through plant transpiration.

The "Kluge-Lüttge Kammer": a preliminary evaluation of an enclosed, Crassulacean acid metabolism (CAM) Mesocosm that allows separation of synchronized and desynchronized contributions of plants to whole system gas exchange.

Crassulacean acid metabolism (CAM) is recognized as a photosynthetic adaptation of plants to arid habitats. This paper presents a proof-of-concept evaluation of partitioning net CO2 exchanges for soil and plants in an arid, exclusively CAM mesocosm, with soil depth and succulent plant biomass approximating that of natural Sonoran Desert ecosystems. We present the first evidence that an enclosed CAM-dominated soil and plant community exposed to a substantial day/night temperature difference (30/20 degrees C), exhibits a diel gas exchange pattern consisting of four consecutive phases with a distinct nocturnal CO2 uptake. These phases were modulated by plant assimilation and soil respiration processes. Day-time stomatal closure of the CAM cycle during phase III was used to eliminate aboveground photosynthetic assimilation and respiration and thereby to estimate belowground plant plus soil respiration. Rapid changes in temperature appeared to synchronize single plant gas exchange but individual plant gas exchange patterns were desynchronized at constant day/night temperatures (25 degrees C), masking the distinct mesocosm pattern. Overall, the mean carbon budget of this CAM model Sonoran Desert system was negative, releasing an average of 22.5 mmol CO2 m-2 d-1. The capacity for nocturnal CO2 assimilation in this exclusively CAM mesocosm was inadequate to recycle CO2 released by plant and soil respiration.

Stochastic noise interferes coherently with a model biological clock and produces specific dynamic behaviour.

The influence of noise is unavoidable in all living systems. Its impact on a model of a biological clock, normally running in regular oscillating modes, is examined. It is shown that in a specific system in which endogenous rhythmicity is produced by a beat oscillator acting on a feedback coupled metabolic pool system, noise can act coherently to produce unexpected dynamic behaviour, running from regular over pseudo-regular to irregular time-structures. If the biological system consists of a set of identical weakly coupled cells, stochasticity may lead to phase decoupling producing irregular spatio-temporal patterns. Synchronization via phase resetting can be achieved by external short-time temperature pulses. Explicit results are obtained for the well-studied circadian photosynthesis oscillations in plants performing crassulacean acid metabolism. Because of the generic structure of the underlying nonlinear dynamics they can, however, be regarded as a general property of the influence of noise on nonlinear excitable systems with fixed points occuring close to limit cycles.

Transitions in photosynthetic parameters of midvein and interveinal regions of leaves and their importance during leaf growth and development.

The areal development of photosynthetic efficiency and growth patterns in expanding leaves of two different dicotyledonous species - Coccoloba uvifera and Sanchezia nobilis - was investigated by imaging both processes repeatedly over 32 days. Measurements were performed using combined imaging systems for chlorophyll fluorescence and growth, with the same spatial resolution. Significant differences in potential quantum yield of photosynthesis (F (v)/F (m)), a parameter indicating the functional status of photosystem II, were found between midvein and interveinal tissue. Although base-tip gradients and spatial patchiness were observed in the distribution of relative growth rate, neither midvein nor interveinal tissue showed such patterns in F (v)/F (m). In young leaves, F (v)/F (m) of the midvein was higher than F (v)/F (m) of interveinal tissue. This difference declined gradually with time, and upon cessation of growth, F (v)/F (m) of interveinal regions exceeded those of midvein tissue. Images of chlorophyll fluorescence quenching showed that DeltaF/F (m)' in the different tissues correlated with F (v)/F (m), indicating that, in these uniformly illuminated leaves, transitions in photosynthetic electron transport activity follow those of predawn quantum efficiency. We explore the implications of these observations during leaf development, discuss effects of sucrose delivery from veins to interveinal areas on relative rates of photosynthetic development in these tissues, and propose that the initially higher photosynthetic activity in the midvein compared to the intervein tissues may supply carbohydrates and energy for leaf growth processes.

Models of fluorescence and photosynthesis for interpreting measurements of solar-induced chlorophyll fluorescence.

We have extended a conventional photosynthesis model to simulate field and laboratory measurements of chlorophyll fluorescence at the leaf scale. The fluorescence paramaterization is based on a close nonlinear relationship between the relative light saturation of photosynthesis and nonradiative energy dissipation in plants of different species. This relationship diverged only among examined data sets under stressed (strongly light saturated) conditions, possibly caused by differences in xanthophyll pigment concentrations. The relationship was quantified after analyzing data sets of pulse amplitude modulated measurements of chlorophyll fluorescence and gas exchange of leaves of different species exposed to different levels of light, CO2, temperature, nitrogen fertilization treatments, and drought. We used this relationship in a photosynthesis model. The coupled model enabled us to quantify the relationships between steady state chlorophyll fluorescence yield, electron transport rate, and photosynthesis in leaves under different environmental conditions. KEY POINTS: Light saturation of photosynthesis determines quenching of leaf fluorescenceWe incorporated steady state leaf fluorescence in a photosynthesis model.

Functional dynamics of plant growth and photosynthesis--from steady-state to dynamics--from homogeneity to heterogeneity.

Plants are much more dynamic than we usually expect them to be. This dynamic behaviour is of paramount importance for their performance under natural conditions, when resources are distributed heterogeneously in space and time. However, plants are not only the cue ball of their physical and chemical environment. Endogenous rhythms and networks controlling photosynthesis and growth buffer plant processes from external fluctuations. This review highlights recent evidence of the importance of dynamic temporal and spatial organization of photosynthesis and of growth in leaves and roots. These central processes for plant performance differ strongly in their dependence on environmental impact and endogenous properties, respectively. Growth involves a wealth of processes ranging from the supply of resources from external and internal sources to the growth processes themselves. In contrast, photosynthesis can only take place when light and CO2 are present and thus clearly requires 'input from the environment'. Nevertheless, growth and photosynthesis are connected to each other via mechanisms that are still not fully understood. Recent advances in imaging technology have provided new insights into the dynamics of plant-environment interactions. Such processes do not only play a crucial role in understanding stress response of plants under extreme environmental conditions. Dynamics of plants under modest growth conditions rise from endogenous mechanisms as well as exogenous impact too. It is thus an important task for future research to identify how dynamic external conditions interact with plant-internal signalling networks to optimize plant behaviour in real time and to understand how plants have adapted to characteristic spatial and temporal properties of the resources from their environment, on which they depend on.

Period-2 cycles and 2:1 phase locking in a biological clock driven by temperature pulses.

Crassulacean acid metabolism (CAM) serves as a botanical model system for the investigation of circadian rhythmicity. In a new set of experiments with the obligatory CAM plant Kalanchoë daigremontiana the response to periodic stimulations with temperature pulses has been studied. On the basis of an experimental phase-response curve of net CO(2)-gas exchange the effect of periodic stimulation has been simulated using a finite-difference equation. These simulations revealed the locations of two period-2 cycles in the CO(2) uptake of the CAM plant. In subsequent experiments based upon the simulated bifurcation diagram the position and amplitude of one of these cycles were confirmed, while experimental evidence for the second cycle could be found. Possible roles of such dynamics for the functioning of the biological clock are discussed.

Spatiotemporal variation of metabolism in a plant circadian rhythm: the biological clock as an assembly of coupled individual oscillators.

The complex dynamic properties of biological timing in organisms remain a central enigma in biology despite the increasingly precise genetic characterization of oscillating units and their components. Although attempts to obtain the time constants from oscillations of gene activity and biochemical units have led to substantial progress, we are still far from a full molecular understanding of endogenous rhythmicity and the physiological manifestations of biological clocks. Applications of nonlinear dynamics have revolutionized thinking in physics and in biomedical and life sciences research, and spatiotemporal considerations are now advancing our understanding of development and rhythmicity. Here we show that the well known circadian rhythm of a metabolic cycle in a higher plant, namely the crassulacean acid metabolism mode of photosynthesis, is expressed as dynamic patterns of independently initiated variations in photosynthetic efficiency (phi(PSII)) over a single leaf. Noninvasive highly sensitive chlorophyll fluorescence imaging reveals randomly initiated patches of varying phi(PSII) that are propagated within minutes to hours in wave fronts, forming dynamically expanding and contracting clusters and clearly dephased regions of phi(PSII). Thus, this biological clock is a spatiotemporal product of many weakly coupled individual oscillators, defined by the metabolic constraints of crassulacean acid metabolism. The oscillators operate independently in space and time as a consequence of the dynamics of metabolic pools and limitations of CO(2) diffusion between tightly packed cells.