Staff Publications

Staff Publications

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    'Staff publications' is the digital repository of Wageningen University & Research

    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

    Publications authored by the staff of the Research Institutes are available from 1995 onwards.

    Full text documents are added when available. The database is updated daily and currently holds about 240,000 items, of which 72,000 in open access.

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Leaf phenolics and seaweed tannins : analysis, enzymatic oxidation and non-covalent protein binding
Vissers, Anne M. - \ 2017
University. Promotor(en): Harry Gruppen; Wouter Hendriks, co-promotor(en): Jean-Paul Vincken. - Wageningen : Wageningen University - ISBN 9789463432023 - 154
phenols - leaves - seaweeds - tannins - beta vulgaris - laminaria - proteins - catechol oxidase - nuclear magnetic resonance spectroscopy - in vitro - mass spectrometry - browning - fermentation - animal feeding - fenolen - bladeren - zeewieren - tanninen - eiwitten - kernmagnetische resonantiespectroscopie - massaspectrometrie - bruinkleuring - fermentatie - diervoedering

Upon extraction of proteins from sugar beet leaves (Beta vulgaris L.) and oarweed (Laminaria digitata) for animal food and feed purposes, endogenous phenolics and proteins can interact with each other, which might affect the protein’s applicability. Sugar beet leaf proteins might become covalently modified by phenolics through polyphenol oxidase (PPO) activity. Oligomeric phenolics from seaweed (so-called phlorotannins (PhT)) might bind non-covalently to protein. The first aim of this thesis was to study factors involved in protein modification by phenolics. The second aim was to investigate the effect of PhT supplementation to feed on in vitro ruminal fermentation.

Besides PPO activity and the amount of low molecular weight phenolic substrates present, brown colour formation in sugar beet leaves was dependent on the amount of phenolics, which do not serve as a substrate of PPO. These non-substrate phenolics can engage in browning reactions by oxidative coupling and subsequent coupled oxidation of the products formed. Similar reactions might also be involved in covalent protein modification by phenolics, and therewith protein properties.
High molecular weight PhT from L. digitata could potentially modify protein properties by non‑covalent interactions. L. digitata contained PhT with subunits mainly connected via C‑O-C linkages, as determined using NMR spectroscopy. Further mass spectrometric analysis revealed the presence of a wide range of oligomers with degrees of polymerisation between 3 and 27. The interaction between PhT and proteins (b-casein and bovine serum albumin) was studied using model systems with different pH values, representing the various environments throughout the ruminants digestive tract. Phlorotannins bound to protein independent of pH, and broadened the pH range of protein precipitation from 0.5 to ~1.5 pH unit around the protein’s pI. At the pH of the abomasum of 2-3, the proteins re-solubilised again, presumably by increase in their net charge. Due to their ability to form water insoluble complexes, PhT could improve ruminal fermentation in vitro in a dose dependent manner, resulting in lower methane production and ammonia (NH3) concentration. The decreased NH3 concentration reflected decreased dietary protein breakdown in the rumen, which is considered a nutritional and environmental benefit.

Sugar beet leaves for functional ingredients
Tamayo Tenorio, Angelica - \ 2017
University. Promotor(en): Remko Boom, co-promotor(en): Atze Jan van der Goot. - Wageningen : Wageningen University - ISBN 9789463431378 - 188
sugarbeet - leaves - thylakoids - cellulosic fibres - food - surface proteins - food crops - protein extraction - suikerbieten - bladeren - thylakoïden - cellulosevezels - voedsel - oppervlakte-eiwitten - voedselgewassen - eiwitextractie

Plant leaves are recognised as a potential source for food applications based on their nutritional profile and interesting technological properties of leaf components, and based on the large availability of plant leaves in agricultural waste streams. Besides proteins, leaves have a rich nutritional profile (e.g. dietary fibres, minerals and secondary metabolites) and consist of complex biological structures (e.g. chloroplastic membranes) that can be explored as novel fractions that ultimately broaden the use of leaves. The overall aim of this thesis is to explore green leaves as a food source, with emphasis on neglected leaf fractions. This thesis describes a processing approach that aims at separating/generating enriched- functional fractions rather than pure components, and highlights the implications for value creation out of green leaves. The extraction of leaf membrane proteins is investigated using a proteomics extraction method, while the properties of other valuable leaf components (complexes and fibres) are analysed for techno-functional applications. Furthermore, the feasibility of leaves as a food source is studied at an industrial scale, considering large scale processing and options for leaf stabilisation.

The extraction of proteins from sugar beet leaves is evaluated in Chapter 2 by using a traditional heat coagulation method. The heat treatment is thought to precipitate the insoluble proteins together with fibres, chlorophyll and other components, resulting in a green curd. Therefore, the distribution of soluble and insoluble proteins was followed along the extraction process to discern the effect of the heating step on protein fractionation. This study showed that both soluble and insoluble protein distribute almost evenly over the leaf fractions juice, pulp, supernatant and final pellet. The even distribution of the proteins was attributed to the anatomy of leaves and their biological function, which is predominantly the enzymatic activity related to photosynthesis instead of protein storage, which occurs in other plant tissues. This chapter further concludes that striving for high purity severely compromises the yield, and consequently results in inefficient use of the leave proteins.

Chapter 3 describes the application of proteomic analytical extraction protocols to analyse the fractionation behaviour of leaf proteins. This analysis lead to the translation into food- grade processes based on four fundamental extraction steps: (1) tissue disruption, (2) enzymatic inhibition, (3) removal of interfering compounds, and (4) protein fractionation and purification. Part of these extraction steps can be translated into food-grade alternatives, while the processing conditions determine the potential properties for food of the final products. Nevertheless, it was concluded that harsh and/or non-food grade conditions were required to isolate the leaf membrane proteins with high purity. Those results were explained by the fact that membrane proteins are heterogeneous w.r.t. charge, hydrophobicity, post- translational modification and complexation, leading to non-selective behaviour when compared with a single pool of proteins.

Given the large challenges in isolating membrane proteins from leaves, we studied another approach in which green leaves are considered as a source of naturally structured elements that have relevant techno-functional properties for food products, like the chloroplastic membranes (i.e. thylakoid membranes) and cellulose-rich fibres. Chapter 4 describes the properties of thylakoid membranes and their emulsifying mechanism. These membranes showed surface active properties and their adsorption kinetics were typical for large molecules or soft particles. The thylakoid fragments can effectively stabilise emulsion droplets, even though aggregation was observed already during emulsion preparation and increased with increased thylakoid concentration. Both composition and structure make thylakoid membranes suitable as a biobased material for food and pharma applications.

To continue exploring valuable fractions from leaves, Chapter 5 reports on the interfacial behaviour of cellulose-rich particles obtained from leaf pulp. Cellulosic particles were produced from the pulp obtained after leaf pressing. The particles spontaneous adsorption onto the oil-water interface and interfacial behaviour similar to that of solid particles. Addition of cellulosic particles to oil-in-water emulsions resulted in stable emulsions above a particle concentration of 0.1 w/v%, although phase separation was observed. The particle fines (0.04 – 1.0 µm) stabilised the droplet interface, while large particles formed a network in the continuous phase and rendered a top (green) phase in the emulsions. Finding applications for leaf side streams, like leaf pulp, broadens the options for total leaf processing and contributes to resource use optimisation.

A sustainability assessment of leaf processing is discussed in Chapter 6, considering the challenges that may appear at industrial scale. The seasonal availability of sugar beet plants implies the need of processing large amounts of biomass within a short time due to their high moisture content (85 - 90%) and their sensitivity to spoilage. Processing options were evaluated on their resource use efficiency in terms of energy requirement and exergy indicators. A decentralised process constitutes a good option compared to freezing, since solid side streams can be directly returned the land, leaving nutrients to the soil, and reducing transportation loads. With a decentralised process, freezing of the leaves becomes unnecessary; the leaf juice is transported while chilled, resembling the transportation of fresh milk that is also chill-transported from the farm to a central factory.

Chapter 7 concludes this thesis with a general discussion of the main findings. An integrated process for leaf valorisation is described, which combines the production of functional fractions with the production of bulk products such as protein-rich and fibre-rich fractions. A compilation of data on protein yield and protein purity of fractions obtained from protein crops (e.g. soy, lupine beans, pulses) and from photosynthetic active tissues (e.g., leaves, algae, duckweed) is included. Protein crops reach 50 - 60% protein yield with a protein purity of ~ 90%, whereas leaves and other photosynthetic active tissues reach similar protein purity (60 – 80 w/w% protein) but at much lower yields (10%). We hypothesize that the low yields are due to the small length scale in which protein is structured inside the leaves and the lack of protein storage anatomy in these tissues. Therefore, we conclude that leaf valorisation requires non-conventional approaches that go beyond higher extraction yields but that consider a complete use of the biomass.

Dynamic photosynthesis under a fluctuating environment: a modelling-based analysis
Morales Sierra, Alejandro - \ 2017
University. Promotor(en): Paul Struik; Jaap Molenaar, co-promotor(en): Xinyou Yin; Jeremy Harbinson. - Wageningen : Wageningen University - ISBN 9789463430456 - 282
photosynthesis - modeling - analysis - environmental factors - light - canopy - leaves - crop physiology - metabolism - fotosynthese - modelleren - analyse - milieufactoren - licht - kroondak - bladeren - gewasfysiologie - metabolisme

In their natural environment, leaves are exposed to rapid fluctuations of irradiance. Research on CO2 assimilation under fluctuating irradiance often relies on measurements of gas exchange during transients where irradiance is rapidly increased or decreased, after the leaf has adapted to a particular set of environmental conditions. In the field, such increases and decreases occur mostly because of sunflecks (rapid increases in irradiance on a low irradiance background) created by gaps in the canopy and plant movement by wind, and cloudflecks (rapid decreases in irradiance on a high irradiance background) generated by clouds that transiently block the sun.

In this dissertation, the metabolic regulation of photosynthesis and how this may limit dynamic CO2 assimilation is studied in silico with the development and application of simulation models. In order to support the development of the models, a review of the literature was performed as well as an experiment designed to generate data on dynamic CO2 assimilation for different photosynthetic mutants of Arabidopsis thaliana. In addition to providing these models to the research community, this dissertation also identifies multiple targets that may be used for improving dynamic CO2 assimilation in plants. It further demonstrates that the dynamic responses of CO2 assimilation to changes in irradiance has a significant effect on canopy CO2 assimilation, even for dense canopies exposed to open skies, resembling the conditions of commercial crops.

In Chapter 1, the context of this dissertation is presented. The societal relevance of this research is argued, making reference to the role that photosynthesis could play in addressing global problems such as food and energy security. The necessary background on the physiology of photosynthesis is provided, with special emphasis on the terminology and concepts required to understand the rest of the dissertation, with the aim of making the contents more accessible to a wider audience. Then, prior literature on the specific topics of this dissertation (i.e., photosynthesis in a dynamic environment and its mathematical modelling) is presented, with a chronological approach that analyses the evolution of ideas and methodologies up to the present.

In Chapter 2, the current literature on dynamic CO2 assimilation is reviewed, with an emphasis on the effects of environmental conditions ([CO2], temperature, and air humidity) on the rates of photosynthetic induction and loss of induction. This review reveals major knowledge gaps, especially on the loss of induction. The little data available indicates that rates of photosynthetic induction increase with [CO2], which could be explained by a weak effect on Rubisco activation and a strong effect on stomatal opening. Increases in temperature also increase the rates of photosynthetic induction, up to an optimum, beyond which a strong negative effect can be observed, which could be attributed to deactivation of Rubisco activase.

In Chapter 3, an experiment is presented that makes use of several photosynthetic mutants of A. thaliana. Downregulating non-photochemical quenching and sucrose synthesis did not have any significant effect on dynamic CO2 assimilation, whereas CO2 diffusion and Rubisco activation exerted stronger limitations. Further analysis reveals that whether stomatal opening limits CO2 assimilation after an increase in irradiance depends on the stomatal conductance prior to the change in irradiance. A threshold value of 0.12 mol m−2 s−1 (defined for fluxes of water vapour) could be defined, above which stomata did not affect the rates of photosynthetic induction. The comparison of measurements across irradiance levels also indicated that the apparent rate constant of Rubisco activation is irradiance-dependent, at least for irradiance levels below 150 μmol m−2 s−1.

In Chapter 4, a phenomenological model of leaf-level CO2 assimilation is presented. The model is described in detail and all the parameters are first estimated with published data, and later refined by fitting the model to the data from Chapter 3. Additional data from the experiment in Chapter 3 is used to validate predictions of CO2 assimilation under lightflecks for the different photosynthetic mutants. The model predicts accurately dynamic CO2 assimilation for the different photosynthetic mutants by only modifying those parameters that are affected by the mutation. This demonstrates that the model has a high predictive power and that the equations, although phenomenological in nature, have a solid physiological basis.

The model is further used to analyse, in silico, the limitations imposed by different photosynthetic processes on dynamic CO2 assimilation at the leaf and canopy level, allowing a more in depth analysis than in Chapter 3. The analysis demonstrates that results obtained at the leaf level should not be extrapolated directly to the canopy level, as the spatial and temporal distribution of irradiance within a canopy is more complex than what is achieved in experimental protocols. Both at the leaf and canopy level, CO2 diffusion is strongly limiting, followed by photoinhibition, chloroplast movements and Rubisco activation.

In Chapter 5, a mechanistic model of the dynamic, metabolic regulation of the electron transport chain is presented. The model is described in detail and all the parameters are estimated from published literature, using measurements on A. thaliana when available. Predictions of the model are tested with steady-state and dynamic measurements of gas exchange, chlorophyll fluorescence and absorbance spectroscopy on A. thaliana, with success.

The analysis in silico indicates that a significant amount of alternative electron transport is required to couple ATP and NADPH production and demand, and most of it is associated with nitrogen assimilation and export of redox power through the malate shuttle. The analysis also reveals that the relationship between ATP synthesis and the proton motive force is highly regulated by the concentrations of substrates (ADP, ATP and inorganic phosphate), and this regulation facilitates an increase in non-photochemical quenching under conditions of low metabolic activity in the stroma.

In Chapter 6, the findings of Chapters 2–5 are summarised and employed to answer in detail the four research questions formulated in Chapter 1. Of great interest is the identification of six potential targets that may be used to improve dynamic CO2 assimilation. These targets are: (i) regulation of Rubisco activity through changes in the amount or regulation of Rubisco activase, (ii) acceleration of stomatal opening and closure, (iii) a lower /ATP for ATP synthesis, (iv) faster relaxation of non-photochemical quenching, (v) reduced chloroplast movements, and (vi) reduced photoinhibition by increased rates of repair of Photosystem II.

Super-performance in a palm species
Jansen, Merel - \ 2016
University. Promotor(en): Niels Anten; Pieter Zuidema, co-promotor(en): Frans Bongers; M. Martínez-Ramos. - Wageningen : Wageningen University - ISBN 9789462579996 - 193
chamaedorea elegans - understorey - tropical forests - spatial variation - leaves - growth - population ecology - defoliation - genetic variation - onderlaag - tropische bossen - ruimtelijke variatie - bladeren - groei - populatie-ecologie - ontbladering - genetische variatie

The world is changing rapidly due to anthropogenic disturbance. Effects include: global warming, massive pollution, a changed global nitrogen cycle, high rates of land-use change, and exotic species spread. This has a tremendous impact on both natural and agricultural systems. To understand these impacts, good understanding of ecological systems and underlying drivers is necessary. Ecological systems can be studied at different levels of aggregation. Different levels of aggregation influence each other and are also influenced by external drivers like the environment. The population level is of particular interest, because many important ecological processes occur at the population level, like evolution, extinction, and invasion. Ecologists are increasingly recognizing that population processes are strongly influenced by one level of aggregation lower, the individual level. Individual heterogeneity (i.e. differences between individuals in performance), determines many population processes including population growth rate. However, the exact relations between individual heterogeneity, the external drivers of it, and the population level are not always well understood. Furthermore, methods to analyze these relations are not always available.

Individual heterogeneity occurs at different temporal scales, ranging from short- to long-term performance differences between individuals, where short- and long-term refer to the expected lifespan of the species in question. Short-term differences between individuals are relatively easily identifiable and are common in almost all species. But long-term differences are much harder to determine especially for long-lived organisms. Long-term differences between individuals in reproduction have been identified for several animal species, and in growth for several tree species, but less is known about the existence of such differences in other life forms (e.g. palms, lianas or clonal plants). Quantifying the extent to which individuals differ is essential for understanding the influence of individual heterogeneity on population processes. Super-performing individuals (i.e. individuals that persistently grow faster and reproduce more than others), probably contribute more to the growth of the population and therefore to future generations. Future populations will, therefore, have the genetic characteristics of the super-performers. Which characteristics this will be, depends on the genetic and environmental drivers of super-performance. Full understanding of the influence of individual heterogeneity on population processes, therefore, requires knowledge of the underlying causes of individual heterogeneity.

For many species, it is known that spatial variation in environmental conditions can cause short-term performance differences between individuals, but it is often not clear if the same environmental factors that cause short-term performance differences are also the environmental factors that cause long-term performance differences. Furthermore, genetic variation is known to cause performance differences, but to what extent is not well studied in natural long-lived plant populations. Within-population genetic variation can be maintained in habitats that are characterized by strong temporal or spatial heterogeneity in environmental conditions if the performance of a genotype relative to others depends on the environment it experiences.

Super-performing individuals possibly play an important role in the resistance and resilience of populations to disturbance (i.e. maintaining and recovering population growth rate under stress), because super-performers potentially contribute more to the recovery of the population. However, this depends on the relative tolerance to disturbance of super-performers compared to under-performers. A positive relation between performance and tolerance would make super-performers more important, while a negative relation would make them less important. Many types of disturbances entail leaf loss and tolerance to leaf loss is associated with performance being larger than what one would assume based on the amount of leaf area loss. Tolerance can be achieved by compensating for leaf loss in terms of growth rate, which entails either allocating more new assimilates to leaves, allocating new assimilates more efficiently to leaf area (i.e. by increasing specific leaf area), or growing faster with existing leaf area (i.e. by increasing net assimilation rate). Genetic variation in tolerance and compensatory responses would allow populations to adapt to changes in disturbance events that entail leaf loss.

Individual heterogeneity could also have implications for management. Plant and animal populations are managed at many different levels ranging from harvest from natural populations to modern agricultural practices. When harvesting from natural populations, it might be beneficial to spare the individuals that are most important for future production. Individuals could be spared, either because they contribute most to population growth, because they are tolerant to harvesting (which is relevant when only part of a plant is harvested), or when they start producing less or lower quality product. The productivity of natural populations could also be increased by actively promoting those environmental conditions and genotypes that allow for high productivity, which is the basis of agriculture and common practice in forest management. To determine how this can best be done, knowledge of the causes of individual heterogeneity is necessary.

The general aim of this thesis is to identify and quantify the mechanisms that determine individual heterogeneity and to determine how this heterogeneity, in turn, affects population level processes. This aim was divided into four main questions that I addressed: (1) To what extent do individuals differ in performance? (2) What causes individual heterogeneity in performance? (3) What are the demographic consequences of individual heterogeneity? (4) Can individual differences be used to improve the management of populations? To answer these questions, we used the tropical forest understorey palm Chamaedorea elegans as a study system, of which the leaves are an important non-timber forest product that is being used in the floral industry worldwide. We collected demographic data, measured spatial variation in environmental conditions, and applied a defoliation treatment to simulate leaf harvesting, in a natural population in Chiapas, Mexico. Furthermore, we grew seedlings from different mothers from our study population in the greenhouses of Wageningen University, where we also applied a defoliation treatment.

In Chapter 2 we quantified the extent to which individuals differ in long-term growth rate, and analyzed the importance of fast growers for population growth. We reconstructed growth histories from internodes and showed that growth differences between individuals are very large and persistent in our study population. This led to large variation in life growth trajectories, with individuals of the same age varying strongly in size. This shows that not only in canopy trees but also in species in the light limited understorey growth differences can be very large. Past growth rate was found to be a very good predictor of current performance (i.e. growth and reproduction). Using an Integral Projection Model (i.e. a type of demographic model) that was based on size and past growth rate, we showed that fast-growing individuals are much more important for population growth than others: the 50% fastest growing individuals contributed almost two times as much to population growth as the 50% slowest growing individuals.

In Chapter 3 we analyzed the extent to which observed long-term growth differences can be caused by environmental heterogeneity. Short-term variation in performance was mainly driven by light availability, while soil variables and leaf damage had smaller effects, and spatial heterogeneity in light availability and soil pH were autocorrelated over time. Using individual-based simulation models, we analyzed the extent to which spatial environmental heterogeneity could explain observed long-term variation in growth, and showed that this could largely be explained if the temporal persistence of light availability and soil pH was taken into account. We also estimated long-term inter-individual variation in reproduction to be very large. We further analyzed the importance of temporal persistence in environmental variation for long-term performance differences, by analyzing the whole range of values of environmental persistence, and the strength of the effect of the environmental heterogeneity on short-term performance. We showed that long-term performance differences become large when either the strength of the effect of the environmental factor on short-term performance is large, or when the spatial variation in the environmental factor is persistent over time. This shows that an environmental factor that in a short-term study might have been dismissed as unimportant for long-term performance variation, might, in reality, contribute strongly.

In Chapter 4 we tested for genetic variation in growth potential, tolerance to leaf loss, compensatory growth responses, and if growth potential and tolerance were genetically correlated in our study population. We quantified compensatory responses with an iterative growth model that takes into account the timing of leaf loss. Genetic variation in growth potential was large, and plants compensated strongly for leaf loss, but genetic variation in tolerance and compensatory growth responses was very limited. Growth performances in defoliated and undefoliated conditions were positively genetically correlated (i.e. the same genotypes perform relatively well compared to others, both with and without the stress of leaf loss). The high genetic variation in growth potential and the positive correlation between treatments suggests that the existence of super-performing individuals in our study population likely has (at least in part) a genetic basis. These super-performing individuals, that grow fast even under the stress of leaf loss, possibly contribute disproportionately to population resistance and resilience to disturbance. The low genetic variation in tolerance and compensatory responses, however, suggests that populations might have limited ability to adapt to changes in disturbance regimes that entail increases in leaf loss. Furthermore, the high genetic variation in growth potential could potentially be used in management practices like enrichment planting.

In Chapter 5 we explore the potential of using individual heterogeneity to design smarter harvest schemes, by sparing individuals that contribute most to future productivity. We tested if fast and slow growers, and small and large individuals, responded differently to leaf loss in terms of vital rates, but found only very limited evidence for this. Using Integral Projection Models that were based on stem length and past growth rate, we simulated leaf harvest over a period of 20 years, in several scenarios of sparing individuals, which we compared to “Business as usual” (i.e. no individuals being spared, BAU). Sparing individuals that are most important for population growth, was beneficial for population size (and could, therefore, reduce extinction risk), increased annual leaf harvest at the end of the simulation period, but cumulated leaf harvest over 20 years was much lower compared to BAU. Sparing individuals that produced leaves of non-commercial size (i.e. <25cm), therefore allowing them to recover, also resulted in a lower total leaf harvest over 20 years. However, a much higher harvest (a three-fold increase) was found when only leaves of commercial size were considered. These results show that it is possible to increase yield quality and sustainability (in terms of population size) of harvesting practices, by making use of individual heterogeneity. The analytical and modeling methods that we present are applicable to any natural system from which either whole individuals, or parts of individuals, are harvested, and provide an extra tool that could be considered by managers and harvest practitioners to optimize harvest practices.

In conclusion, in this thesis, I showed that in a long-lived understorey palm growth differences are very large and persistent (Chapter 2) and that it is likely that long-term differences in reproduction are also very large (Chapter 3). I also showed that spatial heterogeneity in environmental conditions can to a large extent explain these differences and that when evaluating the environmental drivers of individual heterogeneity, it is important to take the persistence of spatial variation into account (Chapter 3). Individual heterogeneity also is partly genetically determined. I showed that genetic variation in growth potential to be large (Chapter 4), and that fast growers keep on growing fast under the stress of leaf loss (Chapters 4,5). Therefore it is likely that genetic variation contributes to long-term differences between individuals. Genetic variation for tolerance and compensatory responses was estimated to be low (Chapter 4), suggesting that the adaptive potential of our study population to changes in disturbance events that entail leaf loss might be low. I also showed that super-performing individuals are much more important for the growth of the population (Chapter 2) and that individuals that are important for future production could be used to improve the management of natural populations (Chapter 5).

This study provides improved insight into the extent of individual heterogeneity in a long-lived plant species and its environmental and genetic drivers, and clearly shows the importance of individual heterogeneity and its drivers for population processes and management practices. It also presents methods on how persistent performance differences between individuals can be incorporated into demographic tools, how these can be used to analyze individual contributions to population dynamics, to extrapolate short-term to long–term environmental effects, and to analyze smart harvesting scenarios that take differences between individuals into account. These results indicate that individual heterogeneity, underlying environmental and genetic drivers, and population processes are all related. Therefore, when evaluating the effect of environmental change on population processes, and in the design of management schemes, it is important to keep these relations in mind. The methodological tools that we presented provide a means of doing this.

Arriving at the right time : a temporal perspective on above-belowground herbivore interactions
Wang, Minggang - \ 2016
University. Promotor(en): Wim van der Putten, co-promotor(en): T.M. Bezemer; A. Biere. - Wageningen : Wageningen University - ISBN 9789462578142 - 174 p.
herbivores - aboveground belowground interactions - herbivory - defence mechanisms - roots - leaves - mycorrhizas - population dynamics - soil biology - herbivoren - boven- en ondergrondse interacties - herbivorie - verdedigingsmechanismen - wortels - bladeren - mycorrhizae - populatiedynamica - bodembiologie
Leaf anatomy and photosynthesis : unravelling the CO2 diffusion pathway in C3 leaves
Berghuijs, H.N.C. - \ 2016
University; KU Leuven. Promotor(en): Paul Struik; Bart M. Nicolaï, co-promotor(en): Xinyou Yin. - Wageningen : Wageningen University - ISBN 9789462577947 - 286 p.
leaves - plant anatomy - photosynthesis - mesophyll - photorespiration - carbon pathways - solanum lycopersicum - bladeren - plantenanatomie - fotosynthese - bladmoes - fotorespiratie - koolstofpathways

Keywords: CO2 diffusion, C3 photosynthesis, mesophyll conductance, mesophyll resistance, re-assimilation, photorespiration, respiration, tomato

Herman Nicolaas Cornelis Berghuijs (2016). Leaf anatomy and photosynthesis; unravelling the CO2 diffusion pathway in C3 leaves. PhD thesis. Wageningen University, Wageningen, The Netherlands, with summaries in English and Dutch. 286 pages

Optimizing photosynthesis can contribute to improving crop yield, which is necessary to meet the increasing global demand for food, fibre, and bioenergy. One way to optimize photosynthesis in C3 plants is to enhance the efficiency of CO2 transport from the intercellular air space to Rubisco. The drawdown of CO2 between these locations is commonly modelled by Fick's first law of diffusion. This law states that the flux from the air spaces to Rubisco is proportional to the difference in partial pressure between these locations. The proportionality constant is the mesophyll conductance. Its inverse is mesophyll resistance. Mesophyll resistance is a complex trait, which lumps various structural barriers for CO2 transport and processes that add or remove CO2 along the diffusion pathway. In order to better understand how and to what extent these factors affect photosynthesis, it is necessary to find a more mechanistic description of CO2 transport in the mesophyll. The aim of this dissertation is to investigate how leaf anatomical properties and CO2 sources and sinks along the CO2­ diffusion pathway in C3 leaves affect the photosynthetic capacity of these leaves. In this study, Solanum lycopersicum was used as a model organism. In a first approach, we developed a model in which we partitioned mesophyll resistance into two sub-resistances. The model assumed that CO2 produced by respiration and photorespiration was released between the two sub-resistance components. By quantifying these resistances using measured thicknesses, exposed mesophyll and chloroplast surfaces, and assumed diffusive properties, we were able to simulate the effect of various anatomical properties on photosynthesis. A disadvantage of this two-resistance approach is that it assumes either that (photo)respiratory CO2 release takes place in the outer cytosol or that there is no CO2 gradient in the cytosol. Therefore, in a second approach we modelled CO2 transport, production and consumption by use of a reaction-diffusion model. This model is more flexible in terms of determining the location of CO2 sources and sinks. We developed methods to estimate physiological parameters of this model using combined gas exchange and chlorophyll fluorescence measurements on leaves. The results suggest that the rate of respiration depends on the oxygen partial pressure, which is often not considered in previous photosynthesis models. We also presented a method to calculate the fraction of (photo)respiratory CO2 that is re-assimilated. We found that this fraction strongly depends on both environmental factors (CO2, irradiance), the location of mitochondria relative to the chloroplast, stomatal conductance and various physiological parameters. The reaction-diffusion model and associated methods presented in this study provide a more mechanistic framework to describe the CO2 diffusion pathway in C3 leaves. This model could, therefore, contribute to identifying targets to increase mesophyll conductance in future research.

Biorefinery of leafy biomass using green tea residue as a model material
Zhang, C. - \ 2016
University. Promotor(en): Johan Sanders, co-promotor(en): Marieke Bruins. - Wageningen : Wageningen University - ISBN 9789462576902 - 156 p.
biorefinery - biomass conversion - leaves - biomass - green tea - tea - alkaline pulping - pectins - lignocellulose - environmental impact - processes - plant protein - food - biobased economy - bioraffinage - biomassaconversie - bladeren - biomassa - groene thee - thee - alkalische pulpbereiding - pectinen - milieueffect - processen - plantaardig eiwit - voedsel


With the rapidly growing world population and improving living standards, food demand is increased with a simultaneous desire for less human impact on the environment, such that “Twice the food production at half the ecological footprint” could be the EU goal for 2050. In fact, a boost in food demand is mainly required in developing countries, where the farmlands are limited and/or they are of poor quality. Rather than improving crop-production yield, developing biorefinery technology with unused biomass, such as leaves, in developing countries may be the key to fulfil the food demand.

Four major components, protein, pectin, lignin, and (hemi-) cellulose, account for more than 70% of the materials in leaves in almost all species. Among these components, protein and pectin can be used in food and animal feed, and they are key components for supplementing food production. However, the production and application of leaf products is limited for four reasons: unstable raw materials, complex components, rigid plant cell walls, and underdeveloped leaf logistics and economics. The limitations cause low pectin and protein yields, and low cost-efficiency in current extraction technologies, including mechanical milling, chemical extraction (acid and alkaline), solvent extraction, and ammonia protein extraction. Development of an integrated process for multiple products might be a good option for leaf biorefinery, but the compatibilities of these processes were unknown.

The aim of this study was to develop new processes and applications that optimally utilize all components, particularly protein, of leafy biomass in the feed and/or food industry using green tea residues as a starting material. The method should also be applicable to other leafy biomass. The research started from the development of alkaline protein extraction technology as presented in Chapter 2. We found that in alkaline protein extraction, temperature, NaOH amount, and extraction time are the parameters determining protein yield, while pH and volume of extraction liquid are critical parameters for production cost. After optimization, more than 90% of leaf protein could be extracted at a cost of 102€/ton protein by single step alkaline extraction. The extracted protein nutritional value was comparable to soybean meal and this technique can be adapted to various leafy biomass. Main drawback of this technique is the overuse of alkali, generation of salts, and the destruction of key amino acids, such as lysine, during the extraction. We tried to overcome its drawbacks by developing integrated process with a recycle for chemicals.

Chapter 3, 4, 5, and 6 refer to the integrated biorefinery. For a better design, we investigated how the alkali aided protein extraction (Chapter 3), and proved that alkaline protein extraction was not facilitated by increased solubility or hydrolysis of protein, but positively correlated to leaf tissue disruption. HG pectin, RGII pectin, polyphenols, and organic acids can be extracted before protein. Protein extraction can then be followed by the extraction of cellulose and hemi-cellulose. RGI pectin and lignin yield were both linearly correlated to protein yield, which indicated that they are likely to be the key limitation to leaf protein extraction. Based on the above findings, an integrated biorefinery that combined protein extraction with a pre-treatment was proposed. In Chapter 4, ethanol, viscozyme, and H2O2 were selected for pre-treatments targeting on the removal of polyphenols and pigments, carbohydrates, and lignin accordingly. Ethanol and viscozyme could extract their targeting components efficiently while H2O2 could bleach GTR with no lignin extracted. The best pre-treatment was the combination of viscozyme and 50% ethanol extraction, which not only reduced the use of alkali by 50%, but also improved protein content and its nutritional value. As pectin can be applied for food or chemicals, enzyme and PBS buffer were investigated for pectin extraction (Chapter 5). Both enzyme and PBS buffer extraction could not only extract high yield HG pectin (predominated by galacturonic acid) with no protein extraction, but also reduced alkali usage in subsequent protein extraction. Pectin obtained using PBS buffer could be present in its native form, which can be precipitated by 40% ethanol. Buffer is suggested to extract pectins when pectins are to be used in food. Otherwise, hydrolyzed pectin that mainly contains galacturonic acid, can be converted to other useful chemicals. For this the enzymatic methods, such as using Viscozyme® L, are recommended.

Alkali usage was further optimized. It was found that by using potassium hydroxide, the protein extraction efficiency was similar to that using sodium hydroxide. The waste water, mainly containing potassium salts, can then be used as fertilizer. This technique is highly depending on the location of factories, which should be built close to the field. Alternatively, calcium hydroxide can be used. As calcium salts can be precipitated by CO2 and calcium hydroxide can be regenerated through burning of the precipitate, this scheme is sustainable and adaptable to most situations. However, as calcium also precipitated pectin, ployphenols, and even proteins, the protein yield is relatively low. Although a pre-treatment can improve extraction efficiency of calcium hydroxide, economic results suggested that a pre-treatment is not necessary unless the products obtained by pre-treatment have an attractive market value.

In Chapter 7, we extend our knowledge on leaf biorefinery with some additional experiments and literature. Simplified models of leaf tissues and cell walls were proposed and used to explain the mechanism of alkaline protein extraction. The models were also used to explain other mechanisms for protein extraction; mechanical milling, steam explosion, acid, and enzyme aided extraction. The possible improvements of leaf biorefinery economics were illustrated either by reducing production cost, by e.g. using counter current extraction or ultrafiltration, or by upgrading product value by applying protein and pectin in food. The processes recommended in this thesis show an excellent prospective, in which they are applicable to other leaf biomass and suitable for small-scale production.

Mating type and sexual fruiting body of Botrytis elliptica, the causal agent of fire blight in lily
Terhem, R.B. ; Staats, M. ; Kan, J.A.L. van - \ 2015
European Journal of Plant Pathology 142 (2015)3. - ISSN 0929-1873 - p. 615 - 624.
cinerea - leaves - resistance - infection - behavior - system
Botrytis elliptica is a necrotrophic pathogen that specifically infects Lilium species. Previous records show that B. elliptica collected in the field can successfully develop apothecia in vitro, however, there are no formal descriptions of apothecia of B. elliptica. The aim of this study was to analyse the sequence of the mating type loci of B. elliptica and produce apothecia in the laboratory in order to describe their morphology. The sequences of both MAT alleles (MAT1-1 or MAT1-2) of B. elliptica were determined and compared to the sister taxa, Botrytis cinerea and Sclerotinia sclerotiorum. Two strains of each mating type were used in crosses under controlled conditions to produce apothecia. Primordium rupture from sclerotial tissue occurred 74 days after fertilization and a mature apothecium formed within 1 month after rupture. The apothecia are 7 to 12 mm in height with a disk of 3 to 4 mm in diameter and 0.5 to 1 mm in thickness. The apothecial disk is usually umbilicate, depressed to funnel and rounded in shape. The number of apothecia growing on a sclerotium was one to nine. Asci are long, cylindrical with a size of 208¿×¿14 µm, thin walled and bearing eight ascospores. Ascospores are hyaline in colour, ellipsoidal with rounded ends, usually 18 to 24 µm in length and 6 to 10 µm in width (mean 19.5¿×¿8 µm). Ascospores were infectious on lily leaves.
Spatial heterogeneity in stomatal features during leaf elongation: an analysis using Rosa hybrida
Fanourakis, D. ; Heuvelink, E. ; Carvalho, S.M.P. - \ 2015
Functional Plant Biology 42 (2015)8. - ISSN 1445-4408 - p. 737 - 745.
relative air humidity - gas-exchange - elevated co2 - conductance - leaves - size - density - adaptation - ontogeny - growth
Within-leaf heterogeneity in stomatal traits poses a key uncertainty in determining a representative value for the whole leaf. Accounting for this heterogeneity, we studied stomatal initiation on expanding leaves and estimated stomatal conductance (gs) of mature leaves. The entire lamina was evaluated at four percentages of full leaflet elongation (FLE; leaflet length relative to its final length) in Rosa hybrida L. plants grown at 60% relative air humidity (RH), and at 100% FLE following cultivation at elevated (95%) RH. Over 80% of the stomata were initiated between 33 and 67% FLE, whereas stomatal growth mostly occurred afterwards. At 100% FLE, the heterogeneity in stomatal density was the result of uneven stomatal differentiation, while an uneven differentiation of epidermal cells contributed to this variation only at elevated RH. Noticeable within-leaf differences (up to 40%) in gs were calculated at 100% FLE. Avoiding leaflet periphery decreased this heterogeneity. Despite the large promotive effect of elevated RH on stomatal and pore dimensions, the within-leaf variation remained unaffected in all characters, besides pore aperture (and, thus, gs). The noted level of within-leaf variation in stomatal features demands a sampling scheme tailored to the leaf developmental stage, the feature per se and the evaporative demand during growth.
Quantifying the source-sink balance and carbohydrate content in three tomato cultivars
Li, T. ; Heuvelink, E. ; Marcelis, L.F.M. - \ 2015
Frontiers in Plant Science 6 (2015). - ISSN 1664-462X
dry-matter production - leaf photosynthesis - plant-growth - leaves - strength - yield - metabolism - simulation - storage - light
Supplementary lighting is frequently applied in the winter season for crop production in greenhouses. The effect of supplementary lighting on plant growth depends on the balance between assimilate production in source leaves and the overall capacity of the plants to use assimilates. This study aims at quantifying the source-sink balance and carbohydrate content of three tomato cultivars differing in fruit size, and to investigate to what extent the source/sink ratio correlates with the potential fruit size. Cultivars Komeet (large size), Capricia (medium size), and Sunstream (small size, cherry tomato) were grown from 16 August to 21 November, at similar crop management as in commercial practice. Supplementary lighting (High Pressure Sodium lamps, photosynthetic active radiation at 1 m below lamps was 162 mu mol photons m(-2) s(-1); maximum 10 h per day depending on solar irradiance level) was applied from 19 September onward. Source strength was estimated from total plant growth rate using periodic destructive plant harvests in combination with the crop growth model TOMSIM. Sink strength was estimated from potential fruit growth rate which was determined from non-destructively measuring the fruit growth rate at non-limiting assimilate supply, growing only one fruit on each truss. Carbohydrate content in leaves and stems were periodically determined. During the early growth stage, Komeet' and Capricia' showed sink limitation and 'Sunstream' was close to sink limitation. During this stage reproductive organs had hardly formed or were still small and natural irradiance was high (early September) compared to winter months. Subsequently, during the fully fruiting stage all three cultivars were strongly source-limited as indicated by the low source/sink ratio (average source/sink ratio from 50 days after planting onward was 0.17, 0.22, and 0.33 for 'Komeet, 'Capricia,' and 'Sunstream,' respectively). This was further confirmed by the fact that pruning half of the fruits hardly influenced net leaf photosynthesis rates. Carbohydrate content in leaves and stems increased linearly with the source/sink ratio. We conclude that during the early growth stage under high irradiance, tomato plants are sink-limited and that the level of sink limitation differs between cultivars but it is not correlated with their potential fruit size. During the fully fruiting stage tomato plants are source-limited and the extent of source limitation of a cultivar is positively correlated with its potential fruit size.
Phloem flow and sugar transport in Ricinus communis L. is inhibited under anoxic conditions of shoot or roots
Peuke, A.D. ; Gessler, A. ; Trumbore, S. ; Windt, C.W. ; Homan, N. ; Gerkema, E. ; As, H. van - \ 2015
Plant, Cell & Environment 38 (2015)3. - ISSN 0140-7791 - p. 433 - 447.
carbon-isotope composition - mushrooms agaricus-bisporus - distance water transport - organic-matter - membrane-permeability - assimilate transport - plants - leaves - starch - stress
Anoxic conditions should hamper the transport of sugar in the phloem, as this is an active process. The canopy is a carbohydrate source and the roots are carbohydrate sinks.By fumigating the shoot with N2 or flooding the rhizosphere, anoxic conditions in the source or sink, respectively, were induced. Volume flow, velocity, conducting area and stationary water of the phloem were assessed by non-invasive magnetic resonance imaging (MRI) flowmetry. Carbohydrates and d13C in leaves, roots and phloem saps were determined. Following flooding, volume flow and conducting area of the phloem declined and sugar concentrations in leaves and in phloem saps slightly increased. Oligosaccharides appeared in phloem saps and after 3 d, carbon transport was reduced to 77%. Additionally, the xylem flow declined and showed finally no daily rhythm. Anoxia of the shoot resulted within minutes in a reduction of volume flow, conductive area and sucrose in the phloem sap decreased. Sugar transport dropped to below 40% by the end of the N2 treatment. However, volume flow and phloem sap sugar tended to recover during the N2 treatment. Both anoxia treatments hampered sugar transport. The flow velocity remained about constant, although phloem sap sugar concentration changed during treatments. Apparently, stored starch was remobilized under anoxia.
Identification, Quantification, and Sensory Characterization of Steviol Glycosides from Differently Processed Stevia rebaudiana Commercial Extracts
Espinoza, M.I. ; Vincken, J.P. ; Sanders, M.G. ; Castro, C. ; Stieger, M.A. ; Agosin, E. - \ 2014
Journal of Agricultural and Food Chemistry 62 (2014)49. - ISSN 0021-8561 - p. 11797 - 11804.
minor diterpene glycosides - liquid-chromatography - rebaudioside-a - transglycosylation - sweeteners - leaves
Stevia rebaudiana is known for its sweet-tasting ent-kaurene diterpenoid glycosides. Several manufacturing strategies are currently employed to obtain Stevia sweeteners with the lowest possible off-flavors. The chemical composition of four commercial S. rebaudiana extracts, obtained by different technologies, was characterized using UHPLC-ESI-MSn. The composition of one of the ethanol-crystallized extracts (EC2) was entirely rebaudioside A, whereas the enzymatically modified (EM) extract contained the lowest concentration of this compound (2.7 mg/100 mg). The membrane-purified (MP) extract had the highest content of minor natural steviol glycosides (23.7 mg/100 mg total extract) versus an average of 2.4 mg/100 mg total extract for the EC samples. Thirteen trained panelists evaluated sweetness, bitterness, licorice, and metallic attributes of all four extracts. The highest licorice intensity (p = 0.05) was found for MP. Both samples EC1 and EC2, despite their different chemical compositions, showed no significant differences in sensory perception.
Monitoring cashew seedlings during interactions with the fungus Lasiodiplodia theobromae using chlorophyll fluorescence imaging
Muniz, C.R. ; Freire, F.C.O. ; Viana, F.M.P. ; Cardoso, J.E. ; Sousa, C.A.F. ; Guedes, M.I.F. ; Schoor, R. van der; Jalink, H. - \ 2014
Photosynthetica 52 (2014)4. - ISSN 0300-3604 - p. 529 - 537.
plant-pathogen interactions - a fluorescence - photosynthetic responses - leaves - identification - parameters - infection - regions - stress
The chlorophyll (Chl) fluorescence imaging technique was applied to cashew seedlings inoculated with the fungus Lasiodiplodia theobromae to assess any disturbances in the photosynthetic apparatus of the plants before the onset of visual symptoms. Two-month-old cashew plants were inoculated with mycelium of L. theobromae isolate Lt19 or Lt32. Dark-adapted and light-acclimated whole plants or previously labelled, single, mature leaf from each plant were evaluated weekly for Chl fluorescence parameters. From 21 to 28 days, inoculation with both isolates resulted in the significantly lower maximal photochemical quantum yield of PSII (Fv/Fm) than those for control samples, decreasing from values of 0.78 to 0.62. In contrast, the time response of the measured fluorescence transient curve from dark-acclimated plants increased in both whole plants and single mature leaves in inoculated plants compared with controls. The Fv/Fm images clearly exhibited photosynthetic perturbations 14 days after inoculation before any visual symptoms appeared. Additionally, decays in the effective quantum yield of PSII photochemistry and photochemical quenching coefficient were also observed over time. However, nonphotochemical quenching increased during the evaluation period. We conclude that Fv/Fm images are the effective way of detecting early metabolic perturbations in the photosynthetic apparatus of cashew seedlings caused by gummosis in both whole plants and single leaves and could be potentially employed in larger-scale screening systems.
Chrysanthemyl diphosphate synthase operates in planta as a bifunctional enzyme with chrysanthemolsynthase activity
Yang, T. ; Gao, L. ; Hu, H. ; Stoopen, G.M. ; Wang, C. ; Jongsma, M.A. - \ 2014
Journal of Biological Chemistry 289 (2014). - ISSN 0021-9258 - p. 36325 - 36335.
dimethylallyl diphosphate - squalene synthase - monoterpene synthase - terpene synthases - biosynthesis - expression - arabidopsis - metabolism - cloning - leaves
Chrysanthemyl diphosphate synthase (CDS) is the first pathway-specific enzyme inthe biosynthesis of pyrethrins, the most widely used plant-derivedpesticide.CDScatalyzes c1’-2-3 cyclopropanation reactions of two molecules of dimethylallyl diphosphate (DMAPP) to yield chrysanthemyl diphosphate (CPP). Three proteinsare known to catalyzethis cyclopropanation reactionof terpene precursors.Two of them, phytoene and squalenesynthase, arebifunctional enzymes with both prenyltransferase and terpenesynthase activity. CDS, the other member,was reported to perform only the prenyltransferase step.Here, we show thatthe NDXXD catalytic motif of CDS,under lowersubstrate conditions prevalent inplants,also catalyzesthe next step converting CPP into chrysanthemolby hydrolyzing the diphosphatemoiety.The enzymatic hydrolysis reactionfollowed conventional Michaelis-Menten kinetics, withaKM value for CPP of 196 µM.For the chrysanthemol synthase activity, DMAPP competed with CPP as substrate. The DMAPP concentration required for half-maximal activity to produce chrysanthemolwas ~100 µM, and significant substrate inhibition was observed at elevated DMAPP concentrations. The N-terminal peptide of CDS was identified as a plastid targeting peptide. Transgenic tobacco plants overexpressing CDS emitted chrysanthemolat arate of 0.12 –0.16 µg•h-1•g-1FW. We propose that CDS should be renamed a chrysanthemol synthase(CHS)utilizingDMAPP as substrate.
Linking leaf initiation to the aerial environment: when air temperature is not the whole story
Savvides, A. - \ 2014
University. Promotor(en): Leo Marcelis, co-promotor(en): Wim van Ieperen; Anja Dieleman. - Wageningen : Wageningen University - ISBN 9789462571136 - 154
bladeren - plantenontwikkeling - initiatie - knoppen - bladmeristemen - licht - luchttemperatuur - leaves - plant development - initiation - buds - leaf meristems - light - air temperature

The initiation of new leaves, which takes place at the shoot apical meristem, is essential for plant growth and development. Leaf initiation rate (LIR) is very sensitive to meristem temperature. However, in practice meristem temperature is hardly ever monitored and air temperature is often used instead. It can be questioned whether relating LIR solely to air temperature is valid. This thesis aims at linking LIR to the aerial environment in two main horticultural crops: tomato and cucumber. It was shown that meristem temperature often differs from air temperature, depending on other environmental factors (e.g. radiation, humidity and wind speed) and species-specific traits. LIR was solely influenced by meristem temperature even when it largely deviated from air temperature. In addition, LIR was reduced at low light levels. Consequently, air temperature is not the whole story when relating leaf initiation to the environment.

Genetic diversity of nitrogen use efficiency in spinach (Spinacia oleraces L.) cultivars using the Ingestad model on hydroponics
Chan-Navarrete, R. ; Kawai, A. ; Dolstra, O. ; Lammerts Van Bueren, E. ; Linden, C.G. van der - \ 2014
Euphytica 199 (2014)1-2. - ISSN 0014-2336 - p. 155 - 166.
nitrate accumulation - crop plants - growth - vegetables - nutrition - photosynthesis - leaves - yield - acid - allocation
Spinach is a leafy vegetable that requires a high N fertilization to have a satisfactory yield and quality, in part because it has poor nitrogen use efficiency (NUE). Therefore, there is a need to breed for cultivars with an excellent NUE. To this end the genetic diversity for NUE-related traits was studied in a diverse set of commercial cultivars. This set was evaluated in a hydroponic system using the Ingestad model; the system was set at a relative growth rate of 0.14 and 0.18 g g-1 day-1 (low and high N, respectively). Experiments were performed at low and high plant density. Traits monitored for single plants included fresh and dry weight, leaf area, specific leaf area, dry weight ratio between root and shoot, and chlorophyll content. The high density experiment showed more genotypic variation for the observed traits than the low density one. Biomass production was considerably lower at low than at high N. Path analysis revealed that leaf area had the highest direct effect on NUE, while specific leaf area was an important trait determining variation in NUE at low N. Slow and fast growing genotypes were shown to use different strategies to utilize N, and these strategies are expressed differently at high and low N availability. This indicates that improving spinach for NUE is feasible using the analysed genotypes as source material, and different strategies can be targeted for adaptation of spinach cultivars to low N conditions.
Evaluation of the nutritive value of muiumba (Baikiaea plurijuga) seeds: chemical composition, in vitro organic matter digestibility and in vitro gas production
Rodrigues, M.A.M. ; Lourenço, A.L. ; Cone, J.W. ; Nunes, F.M. ; Santos, A.S. ; Cordeiro, J.M.M. ; Guedes, C.M.V. ; Ferreira, L.M.M. - \ 2014
SpringerPlus 3 (2014). - ISSN 2193-1801
nonstarch polysaccharides - production profiles - plant materials - dietary fiber - tree fodder - fermentation - feed - fractions - leaves - fruits
One of the main constraints hindering the increase of animal production in semi-arid regions of Africa is the inadequate supply of nutrients during the dry season. Incorporation of alternative feed resources in ruminant diets during this period could be a viable approach to overcome these limitations. The objective of this study was to evaluate the nutritive value of muiumba (Baikiaea plurijuga) tree seeds as an alternative nutrient source for ruminants. Muiumba seeds were compared to other eight feedstuffs including two cereal grains (corn and oat), two wheat by-products (wheat bran and distilled wheat) and four protein meals (coconut meal, sunflower meal, soybean meal and rapeseed meal) as to its chemical composition, in vitro organic matter digestibility (IVOMD) and in vitro gas production. The moderate crude protein concentrations (145 g/kg DM) of muiumba seeds indicate that this feedstuff could not be used as a protein supplement, contrarily to the majority of multipurpose tree seeds. Although the starch content was scarce (15 g/kg DM), the low neutral detergent fibre (235 g/kg DM), low molecular weight sugar (76.1 g/kg DM) and non-starch polysaccharide (510.5 g/kg DM) contents indicate that this feedstuff has potential feeding value. This was confirmed by the IVOMD (0.770) and by the data provided by the in vitro gas production showing that muiumba seeds had high (P <0.05) maximum gas production and fractional fermentation rates, suggesting that these seeds are characterized by a highly fermentable fraction.
Enhancement of crop photosynthesis by diffuse light: quantifying the contributing factors
Li, T. ; Heuvelink, E. ; Dueck, T.A. ; Janse, J. ; Gort, G. ; Marcelis, L.F.M. - \ 2014
Annals of Botany 114 (2014)1. - ISSN 0305-7364 - p. 145 - 156.
modeling canopy photosynthesis - net ecosystem exchange - structural plant-model - chlorophyll fluorescence - deciduous forest - global radiation - direct component - solar-radiation - leaf-area - leaves
Plants use diffuse light more efficiently than direct light. However, experimental comparisons between diffuse and direct light have been obscured by co-occurring differences in environmental conditions (e.g. light intensity). This study aims to analyse the factors that contribute to an increase in crop photosynthesis in diffuse light and to quantify their relative contribution under different levels of diffuseness at similar light intensities. The hypothesis is that the enhancement of crop photosynthesis in diffuse light results not only from the direct effects of more uniform vertical and horizontal light distribution in the crop canopy, but also from crop physiological and morphological acclimation. Tomato (Solanum lycopersicum) crops were grown in three greenhouse compartments that were covered by glass with different degrees of light diffuseness (0, 45 and 71 % of the direct light being converted into diffuse light) while maintaining similar light transmission. Measurements of horizontal and vertical photosynthetic photon flux density (PPFD) distribution in the crop, leaf photosynthesis light response curves and leaf area index (LAI) were used to quantify each factor's contribution to an increase in crop photosynthesis in diffuse light. In addition, leaf temperature, photoinhibition, and leaf biochemical and anatomical properties were studied. The highest degree of light diffuseness (71 %) increased the calculated crop photosynthesis by 7 center dot 2 %. This effect was mainly attributed to a more uniform horizontal (33 % of the total effect) and vertical PPFD distribution (21 %) in the crop. In addition, plants acclimated to the high level of diffuseness by gaining a higher photosynthetic capacity of leaves in the middle of the crop and a higher LAI, which contributed 23 and 13 %, respectively, to the total increase in crop photosynthesis in diffuse light. Moreover, diffuse light resulted in lower leaf temperatures and less photoinhibition at the top of the canopy when global irradiance was high. Diffuse light enhanced crop photosynthesis. A more uniform horizontal PPFD distribution played the most important role in this enhancement, and a more uniform vertical PPFD distribution and higher leaf photosynthetic capacity contributed more to the enhancement of crop photosynthesis than did higher values of LAI.
Metabolomics as a Potential Chemotaxonomical Tool: Application in the Genus Vernonia Schreb
Martucci, M.E.P. ; Vos, R.C.H. de; Carollo, C.A. - \ 2014
PLoS One 9 (2014)4. - ISSN 1932-6203 - 8 p.
ericoides mart. asteraceae - mass-spectrometry - sesquiterpene lactones - chlorogenic acids - flavonoids - identification - compositae - amygdalina - saponins - leaves
The taxonomic classification of the genus Vernonia Schreb is complex and, as yet, unclear. We here report the use of untargeted metabolomics approaches, followed by multivariate analyses methods and a phytochemical characterization of ten Vernonia species. Metabolic fingerprints were obtained by accurate mass measurements and used to determine the phytochemical similarities and differences between species through multivariate analyses approaches. Principal component analysis based on the relative levels of 528 metabolites, indicated that the ten species could be clustered into four groups. Thereby, V. polyanthes was the only species with presence of flavones chrysoeriol-7-O-glycuronyl, acacetin-7-O-glycuronyl and sesquiterpenes lactones piptocarphin A and piptocarphin B, while glaucolide A was detected in both V. brasiliana and V. polyanthes, separating these species from the two other species of the Vernonanthura group. Species from the Lessingianthus group were unique in showing a positive response in the foam test, suggesting the presence of saponins, which could be confirmed by metabolite annotation. V. rufogrisea showed a great variety of sesquiterpene lactones, placing this species into a separate group. Species within the Chrysolaena group were unique in accumulating clovamide. Our results of LC-MS-based profiling combined with multivariate analyses suggest that metabolomics approaches, such as untargeted LC-MS, may be potentially used as a large-scale chemotaxonomical tool, in addition to classical morphological and cytotaxonomical approaches, in order to facilitate taxonomical classifications.
Storage of intact heads prior to processing limits the shelf-life of fresh-cut Lactuca sativa L.
Witkowska, I.M. ; Woltering, E.J. - \ 2014
Postharvest Biology and Technology 91 (2014). - ISSN 0925-5214 - p. 25 - 31.
growth stage - lollo-rosso - antioxidant capacity - induced senescence - leaf senescence - lettuce - quality - spinach - leaves - vegetables
Harvested lettuce heads are usually transported and stored for some period of time under a variety of conditions prior to processing. During storage, especially under suboptimal conditions, nutritional composition of the harvested produce continues to change. The possible impact of prior storage of the heads on the performance of the fresh-cut product has not been quantified, and was the aim of this study. The experiments were performed with three related genotypes of Lactuca sativa L. (butterhead lettuce): two green varieties and one red variety. The effect of prior storage on quality parameters in the stored whole heads and on subsequent fresh-cut quality performance was investigated. In addition, the effect of prior storage of heads with and without their root system and the application of light during storage were investigated. The changes in visual quality, the levels of energy reserves, and some selected senescence markers, i.e. chlorophyll content and electrolyte leakage were evaluated. Despite the relatively high storage temperature of 12 °C, the intact heads still looked fresh even after 17 days of storage. However, a decline in the soluble sugars, a decrease in chlorophyll, and an increase in electrolyte leakage were observed with advancing storage duration. Prior storage of intact heads greatly decreased the shelf-life of the fresh-cut product prepared from these heads. Storage of rooted heads and the continuous application of light (above the light compensation point) did not alter the effect of prior storage of the heads on the quality of the fresh-cut product
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