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Hyperspectral imaging sensors are promising tools for monitoring crop plants or vegetation in different environments. Information on physiology, architecture or biochemistry of plants can be assessed non-invasively and on different scales. For instance, hyperspectral sensors are implemented for stress detection in plant phenotyping processes or in precision agriculture. Up to date, a variety of non-imaging and imaging hyperspectral sensors is available. The measuring process and the handling of most of these sensors is rather complex. Thus, during the last years the demand for sensors with easy user operability arose. The present study introduces the novel hyperspectral camera Specim IQ from Specim (Oulu, Finland). The Specim IQ is a handheld push broom system with integrated operating system and controls. Basic data handling and data analysis processes, such as pre-processing and classification routines are implemented within the camera software. This study provides an introduction into the measurement pipeline of the Specim IQ as well as a radiometric performance comparison with a well-established hyperspectral imager. Case studies for the detection of powdery mildew on barley at the canopy scale and the spectral characterization of Arabidopsis thaliana mutants grown under stressed and non-stressed conditions are presented., Competing Interests: SPECIM, Spectral Imaging Oy Ltd., provided two prototypes of the Specim IQ, a training and prepared . Technical description of the Specim IQ camera system. Additionally, Specim provided financial support to perform the evaluation of the sensor and payed the APC. Scientists from the Forschungszentrum Jülich and Bonn University designed the scientific test cases, analysed the results, produced the graphs and wrote the text. Representatives of the company were not involved in the design of the comparison with the Specim V10E, the case studies and the writing of the text of the case studies.

Sun-induced fluorescence (SIF) in the far-red region provides a new noninvasive measurement approach that has the potential to quantify dynamic changes in light-use efficiency and gross primary production (GPP). However, the mechanistic link between GPP and SIF is not completely understood. We analyzed the structural and functional factors controlling the emission of SIF at 760 nm (F 760 ) in a Mediterranean grassland manipulated with nutrient addition of nitrogen (N), phosphorous (P) or nitrogen-phosphorous (NP). Using the soil-canopy observation of photosynthesis and energy (SCOPE) model, we investigated how nutrient-induced changes in canopy structure (i.e. changes in plant forms abundance that influence leaf inclination distribution function, LIDF) and functional traits (e.g. N content in dry mass of leaves, N%, Chlorophyll a+b concentration (Cab) and maximum carboxylation capacity (V cmax )) affected the observed linear relationship between F 760 and GPP. We conclude that the addition of nutrients imposed a change in the abundance of different plant forms and biochemistry of the canopy that controls F 760 . Changes in canopy structure mainly control the GPP-F 760 relationship, with a secondary effect of Cab and V cmax . In order to exploit F 760 data to model GPP at the global/regional scale, canopy structural variability, biodiversity and functional traits are important factors that have to be considered., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Passive detection of sun-induced chlorophyll fluorescence (SIF) using spectroscopy has been proposed as a proxy to quantify changes in photochemical efficiency at canopy level under natural light conditions. In this study, we explored the use of imaging spectroscopy to quantify spatio-temporal dynamics of SIF within crop canopies and its sensitivity to track patterns of photosynthetic activity originating from the interaction between vegetation structure and incoming radiation as well as variations in plant function. SIF was retrieved using the Fraunhofer Line Depth (FLD) principle from imaging spectroscopy data acquired at different time scales a few metres above several crop canopies growing under natural illumination. We report the first maps of canopy SIF in high spatial resolution. Changes of SIF were monitored at different time scales ranging from quick variations under induced stress conditions to seasonal dynamics. Natural changes were primarily determined by varying levels and distribution of photosynthetic active radiation (PAR). However, this relationship changed throughout the day demonstrating an additional physiological component modulating spatio-temporal patterns of SIF emission. We successfully used detailed SIF maps to track changes in the canopy's photochemical activity under field conditions, providing a new tool to evaluate complex patterns of photosynthesis within the canopy., (© 2016 John Wiley & Sons Ltd.)

Studies on the function of root traits and the genetic variation in these traits are often conducted under controlled conditions using individual potted plants. Little is known about root growth under field conditions and how root traits are affected by agronomic practices in particular sowing density. We hypothesized that with increasing sowing density, root length density (root length per soil volume, cm cm(-3)) increases in the topsoil as well as specific root length (root length per root dry weight, cm g(-1)) due to greater investment in fine roots. Therefore, we studied two spring barley cultivars at ten different sowing densities (24-340 seeds m(-2)) in 2 consecutive years in a clay loam field in Germany and established sowing density dose-response curves for several root and shoot traits. We took soil cores for measuring roots up to a depth of 60 cm in and between plant rows (inter-row distance 21 cm). Root length density increased with increasing sowing density and was greatest in the plant row in the topsoil (0-10 cm). Greater sowing density increased specific root length partly through greater production of fine roots in the topsoil. Rooting depth (D50) of the major root axes (root diameter class 0.4-1.0 mm) was not affected. Root mass fraction decreased, while stem mass fraction increased with sowing density and over time. Leaf mass fraction was constant over sowing density but greater leaf area was realized through increased specific leaf area. Considering fertilization, we assume that light competition caused plants to grow more shoot mass at the cost of investment into roots, which is partly compensated by increased specific root length and shallow rooting. Increased biomass per area with greater densities suggest that density increases the efficiency of the cropping system, however, declines in harvest index at densities over 230 plants m(-2) suggest that this efficiency did not translate into greater yield. We conclude that plant density is a modifier of root architecture and that root traits and their utility in breeding for greater productivity have to be understood in the context of high sowing densities.

Hyperspectral imaging sensors are valuable tools for plant disease detection and plant phenotyping. Reflectance properties are influenced by plant pathogens and resistance responses, but changes of transmission characteristics of plants are less described. In this study we used simultaneously recorded reflectance and transmittance imaging data of resistant and susceptible barley genotypes that were inoculated with Blumeria graminis f. sp. hordei to evaluate the added value of imaging transmission, reflection and absorption for characterisation of disease development. These datasets were statistically analysed using principal component analysis, and compared with visual and molecular disease estimation. Reflection measurement performed significantly better for early detection of powdery mildew infection, colonies could be detected 2 days before symptoms became visible in RGB images. Transmission data could be used to detect powdery mildew 2 days after symptoms becoming visible in reflection based RGB images. Additionally distinct transmission changes occurred at 580-650nm for pixels containing disease symptoms. It could be shown that the additional information of the transmission data allows for a clearer spatial differentiation and localisation between powdery mildew symptoms and necrotic tissue on the leaf then purely reflectance based data. Thus the information of both measurement approaches are complementary: reflectance based measurements facilitate an early detection, and transmission measurements provide additional information to better understand and quantify the complex spatio-temporal dynamics of plant-pathogen interactions.

Background: Three-dimensional canopies form complex architectures with temporally and spatially changing leaf orientations. Variations in canopy structure are linked to canopy function and they occur within the scope of genetic variability as well as a reaction to environmental factors like light, water and nutrient supply, and stress. An important key measure to characterize these structural properties is the leaf angle distribution, which in turn requires knowledge on the 3-dimensional single leaf surface. Despite a large number of 3-d sensors and methods only a few systems are applicable for fast and routine measurements in plants and natural canopies. A suitable approach is stereo imaging, which combines depth and color information that allows for easy segmentation of green leaf material and the extraction of plant traits, such as leaf angle distribution., Results: We developed a software package, which provides tools for the quantification of leaf surface properties within natural canopies via 3-d reconstruction from stereo images. Our approach includes a semi-automatic selection process of single leaves and different modes of surface characterization via polygon smoothing or surface model fitting. Based on the resulting surface meshes leaf angle statistics are computed on the whole-leaf level or from local derivations. We include a case study to demonstrate the functionality of our software. 48 images of small sugar beet populations (4 varieties) have been analyzed on the base of their leaf angle distribution in order to investigate seasonal, genotypic and fertilization effects on leaf angle distributions. We could show that leaf angle distributions change during the course of the season with all varieties having a comparable development. Additionally, different varieties had different leaf angle orientation that could be separated in principle component analysis. In contrast nitrogen treatment had no effect on leaf angles., Conclusions: We show that a stereo imaging setup together with the appropriate image processing tools is capable of retrieving the geometric leaf surface properties of plants and canopies. Our software package provides whole-leaf statistics but also a local estimation of leaf angles, which may have great potential to better understand and quantify structural canopy traits for guided breeding and optimized crop management.

Background: Combined assessment of leaf reflectance and transmittance is currently limited to spot (point) measurements. This study introduces a tailor-made hyperspectral absorption-reflectance-transmittance imaging (HyperART) system, yielding a non-invasive determination of both reflectance and transmittance of the whole leaf. We addressed its applicability for analysing plant traits, i.e. assessing Cercospora beticola disease severity or leaf chlorophyll content. To test the accuracy of the obtained data, these were compared with reflectance and transmittance measurements of selected leaves acquired by the point spectroradiometer ASD FieldSpec, equipped with the FluoWat device., Results: The working principle of the HyperART system relies on the upward redirection of transmitted and reflected light (range of 400 to 2500 nm) of a plant sample towards two line scanners. By using both the reflectance and transmittance image, an image of leaf absorption can be calculated. The comparison with the dynamically high-resolution ASD FieldSpec data showed good correlation, underlying the accuracy of the HyperART system. Our experiments showed that variation in both leaf chlorophyll content of four different crop species, due to different fertilization regimes during growth, and fungal symptoms on sugar beet leaves could be accurately estimated and monitored. The use of leaf reflectance and transmittance, as well as their sum (by which the non-absorbed radiation is calculated) obtained by the HyperART system gave considerably improved results in classification of Cercospora leaf spot disease and determination of chlorophyll content., Conclusions: The HyperART system offers the possibility for non-invasive and accurate mapping of leaf transmittance and absorption, significantly expanding the applicability of reflectance, based on mapping spectroscopy, in plant sciences. Therefore, the HyperART system may be readily employed for non-invasive determination of the spatio-temporal dynamics of various plant properties.

The photosynthesis of various species or even a single plant varies dramatically in time and space, creating great spatial heterogeneity within a plant canopy. Continuous and spatially explicit monitoring is, therefore, required to assess the dynamic response of plant photosynthesis to the changing environment. This is a very challenging task when using the existing portable field instrumentation. This paper reports on the application of a technique, laser-induced fluorescence transient (LIFT), developed for ground remote measurement of photosynthetic efficiency at a distance of up to 50 m. The LIFT technique was used to monitor the seasonal dynamics of selected leaf groups within inaccessible canopies of deciduous and evergreen tree species. Electron transport rates computed from LIFT measurements varied over the growth period between the different species studied. The LIFT canopy data and light-use efficiency measured under field conditions correlated reasonably well with the single-leaf pulse amplitude-modulated measurements of broadleaf species, but differed significantly in the case of conifer tree species. The LIFT method has proven to be applicable for a remote sensing assessment of photosynthetic parameters on a diurnal and seasonal scale; further investigation is, however, needed to evaluate the influence of complex heterogeneous canopy structures on LIFT-measured chlorophyll fluorescence parameters., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Priority effects occur when species that arrive first in a habitat significantly affect the establishment, growth, or reproduction of species arriving later and thus affect functioning of communities. However, we know little about how the timing of arrival of functionally different species may alter structure and function during assembly. Even less is known about how plant density might interact with initial assembly. In a greenhouse experiment legumes, grasses or forbs were sown a number of weeks before the other two plant functional types were sown (PFT) in combination with a sowing density treatment. Legumes, grasses or non-legume forbs were sown first at three different density levels followed by sowing of the remaining PFTs after three or six-weeks. We found that the order of arrival of different plant functional types had a much stronger influence on aboveground productivity than sowing density or interval between the sowing events. The sowing of legumes before the other PFTs produced the highest aboveground biomass. The larger sowing interval led to higher asymmetric competition, with highest dominance of the PFT sown first. It seems that legumes were better able to get a head-start and be productive before the later groups arrived, but that their traits allowed for better subsequent establishment of non-legume PFTs. Our study indicates that the manipulation of the order of arrival can create priority effects which favour functional groups of plants differently and thus induce different assembly routes and affect community composition and functioning.

Background: Most spectral data for the amphibian integument are limited to the visible spectrum of light and have been collected using point measurements with low spatial resolution. In the present study a dual camera setup consisting of two push broom hyperspectral imaging systems was employed, which produces reflectance images between 400 and 2500 nm with high spectral and spatial resolution and a high dynamic range., Methodology/principal Findings: We briefly introduce the system and document the high efficiency of this technique analyzing exemplarily the spectral reflectivity of the integument of three arboreal anuran species (Litoria caerulea, Agalychnis callidryas and Hyla arborea), all of which appear green to the human eye. The imaging setup generates a high number of spectral bands within seconds and allows non-invasive characterization of spectral characteristics with relatively high working distance. Despite the comparatively uniform coloration, spectral reflectivity between 700 and 1100 nm differed markedly among the species. In contrast to H. arborea, L. caerulea and A. callidryas showed reflection in this range. For all three species, reflectivity above 1100 nm is primarily defined by water absorption. Furthermore, the high resolution allowed examining even small structures such as fingers and toes, which in A. callidryas showed an increased reflectivity in the near infrared part of the spectrum., Conclusion/significance: Hyperspectral imaging was found to be a very useful alternative technique combining the spectral resolution of spectrometric measurements with a higher spatial resolution. In addition, we used Digital Infrared/Red-Edge Photography as new simple method to roughly determine the near infrared reflectivity of frog specimens in field, where hyperspectral imaging is typically difficult.

Early water stress recognition is of great relevance in precision plant breeding and production. Hyperspectral imaging sensors can be a valuable tool for early stress detection with high spatio-temporal resolution. They gather large, high dimensional data cubes posing a significant challenge to data analysis. Classical supervised learning algorithms often fail in applied plant sciences due to their need of labelled datasets, which are difficult to obtain. Therefore, new approaches for unsupervised learning of relevant patterns are needed. We apply for the first time a recent matrix factorisation technique, simplex volume maximisation (SiVM), to hyperspectral data. It is an unsupervised classification approach, optimised for fast computation of massive datasets. It allows calculation of how similar each spectrum is to observed typical spectra. This provides the means to express how likely it is that one plant is suffering from stress. The method was tested for drought stress, applied to potted barley plants in a controlled rain-out shelter experiment and to agricultural corn plots subjected to a two factorial field setup altering water and nutrient availability. Both experiments were conducted on the canopy level. SiVM was significantly better than using a combination of established vegetation indices. In the corn plots, SiVM clearly separated the different treatments, even though the effects on leaf and canopy traits were subtle.

Plants experiencing herbivory suffer indirect costs beyond direct loss of leaf area, but differentially so based on the herbivore involved. We used a combination of chlorophyll fluorescence imaging and gas exchange techniques to quantify photosynthetic performance, the efficiency of photochemistry, and heat dissipation to examine immediate and longer-term physiological responses in the desert perennial Datura wrightii to herbivory by tobacco hornworm, Manduca sexta. Herbivory by colony-reared larvae yielded no significant reduction in carbon assimilation, whereas herbivory by wild larvae induced a fast and spreading down-regulation of photosynthetic efficiency, resulting in significant losses in carbon assimilation in eaten and uneaten leaves. We found both an 89 % reduction in net photosynthetic rates in herbivore-damaged leaves and a whole-plant response (79 % decrease in undamaged leaves from adjacent branches). Consequently, herbivory costs are higher than previously estimated in this well-studied plant-insect interaction. We used chlorophyll fluorescence imaging to elucidate the mechanisms of this down-regulation. Quantum yield decreased up to 70 % in a small concentric band surrounding the feeding area within minutes of the onset of herbivory. Non-photochemical energy dissipation by the plant to avoid permanent damage was elevated near the wound, and increased systematically in distant areas of the leaf away from the wound over subsequent hours. Together, the results underscore not only potential differences between colony-reared and wild-caught herbivores in experimental studies of herbivory but also the benefits of quantifying physiological responses of plants in unattacked leaves.

The interaction of plants with their environment is very dynamic. Studying the underlying processes is important for understanding and modeling plant response to changing environmental conditions. Photosynthesis varies largely between different plants and at different locations within a canopy of a single plant. Thus, continuous and spatially distributed monitoring is necessary to assess the dynamic response of photosynthesis to the environment. Limited scale of observation with portable instrumentation makes it difficult to examine large numbers of plants under different environmental conditions. We report here on the application of a recently developed technique, laser-induced fluorescence transient (LIFT), for continuous remote measurement of photosynthetic efficiency of selected leaves at a distance of up to 50 m. The ability to make continuous, automatic, and remote measurements of photosynthetic efficiency of leaves with the LIFT provides a new approach for studying the interaction of plants with the environment and may become an important tool in phenotyping photosynthetic properties in field applications.

Previous studies of elevated carbon dioxide concentration ([CO(2)]) on crop canopies have found that radiation-use efficiency is increased more than radiation-interception efficiency. It is assumed that increased radiation-use efficiency is due to changes in leaf-level physiology; however, canopy structure can affect radiation-use efficiency if leaves are displayed in a manner that optimizes their physiological capacity, even though the canopy intercepts the same amount of light. In order to determine the contributions of physiology and canopy structure to radiation-use and radiation-interception efficiency, this study relates leaf-level physiology and leaf display to photosynthetic rate of the outer canopy. We used a new imaging approach that delivers three-dimensional maps of the outer canopy during the growing season. The 3D data were used to model leaf orientation and mean photosynthetic electron transport of the outer canopy to show that leaf orientation changes did not contribute to increased radiation-use; i.e. leaves of the outer canopy showed similar diurnal leaf movements and leaf orientation in both treatments. Elevated [CO(2)] resulted in an increased maximum electron transport rate (ETR(max)) of light reactions of photosynthesis. Modeling of canopy light interception showed that stimulated leaf-level electron transport at elevated [CO(2)], and not alterations in leaf orientation, was associated with stimulated radiation-use efficiency and biomass production in elevated [CO(2)]. This study provides proof of concept of methodology to quantify structure-function relationships in combination, allowing a quantitative estimate of the contribution of both effects to canopy energy conversion under elevated [CO(2)].

A new, compact, spatially scanning, open-path 2.7 microm tunable diode laser absorption spectrometer with short absorption path lengths below 10 cm was developed to analyze the spatiotemporal dynamics of one-dimensional (1D) spatial water vapor gradients. This spectrometer, which is based on a room-temperature distributed feedback diode laser, is capable of measuring absolute, calibration-free, line-of-sight averaged, but laterally resolved 1D H(2)O concentration profiles with a minimum fractional optical resolution of 2.1x10(-3) optical density (OD) (2.5x10(-4) OD after a background subtraction procedure), which permits a signal-to-noise-ratio of 407 (3400) at 10,000 parts in 10(6) (ppm)H(2)O, or normalized sensitivities of 2.6 ppm x m (0.32 ppm m) at 0.5 Hz duty cycle. The spectrometer's lateral spatial resolution (governed by the 500 microm sampling beam diameter) was validated by analyzing a well-defined laminar jet of nitrogen gas in humidified air. This scanning setup was then used to (a) quantitatively investigate for what we believe to be the first time the H(2)O boundary layer from 0.7 to 11 mm beneath the stomatous side of a single, undetached plant leaf, and (b) to study the temporal boundary layer dynamics and its dependence on stepwise light stimulation of the photosynthetic system. In addition the 2.7 microm diode laser was carefully characterized in terms of spectral purity, beam profile, as well as quasi-static and dynamic wavelength tuning coefficients.

This study reports evidence that the timing of leaf growth responds to developmental and environmental constraints in Clusia spp. We monitored diel patterns of leaf growth in the facultative C(3)-crassulacean acid metabolism (CAM) species Clusia minor and in the supposedly obligate CAM species Clusia alata using imaging methods and followed diel patterns of CO2 exchange and acidification. Developing leaves of well-watered C. minor showed a C3-like diel pattern of gas exchange and growth, with maximum relative growth rate (RGR) in the early night period. Growth slowed when water was withheld, accompanied by nocturnal CO2 exchange and the diel acid change characteristic of CAM. Maximum leaf RGR shifted from early night to early in the day when water was withheld. In well-watered C. alata, similar changes in the diel pattern of leaf growth occurred with the development of CAM during leaf ontogeny. We hypothesize that the shift in leaf growth cycle that accompanies the switch from C3 photosynthesis to CAM is mainly caused by the primary demand of CAM for substrates for nocturnal CO2 fixation and acid synthesis, thus reducing the availability of carbohydrates for leaf growth at night. Although the shift to leaf growth early in the light is presumably associated with the availability of carbohydrates, source-sink relationships and sustained diurnal acid levels in young leaves of Clusia spp. need further evaluation in relation to growth processes.

Plants are exposed to a temporally and spatially heterogeneous environment, and photosynthesis is well adapted to these fluctuations. Understanding of the complex, non-linear dynamics of photosynthesis in fluctuating light requires novel-modeling approaches that involve not only the primary light and dark biochemical reactions, but also networks of regulatory interactions. This requirement exceeds the capacity of the existing molecular models that are typically reduced to describe a partial process, dynamics of a specific complex or its particular dynamic feature. We propose a concept of comprehensive model that would represent an internally consistent, integral framework combining information on the reduced models that led to its construction. This review explores approaches and tools that exist in engineering, mathematics, and in other domains of biology that can be used to develop a comprehensive model of photosynthesis. Equally important, we investigated techniques by which one can rigorously reduce such a comprehensive model to models of low dimensionality, which preserve dynamic features of interest and, thus, contribute to a better understanding of photosynthesis under natural and thus fluctuating conditions. The web-based platform www.e-photosynthesis.org is introduced as an arena where these concepts and tools are being introduced and tested.

We have developed a laser induced fluorescence transient (LIFT) technique and instrumentation to remotely measure photosynthetic properties in terrestrial vegetation at a distance of up to 50 m. The LIFT method uses a 665 nm laser to project a collimated, 100 mm diameter excitation beam onto leaves of the targeted plant. Fluorescence emission at 690 nm is collected by a 250 mm reflective telescope and processed in real time to calculate the efficiency of photosynthetic light utilization, quantum efficiency of PS II, and the kinetics of photosynthetic electron transport. Operating with peak excitation power of 125 W m-2, and duty cycle of 10-50%, the instrument conforms to laser safety regulations. The LIFT instrument is controlled via an Internet connection, allowing it to operate from remote locations or platforms. Here we describe the theoretical basis of the LIFT methodology, and demonstrate its applications in remote measurements of photosynthetic properties in the canopy of cottonwood and oak trees, and in the rosette of Arabidopsis mutants.