This review presents a comprehensive and systematic study of the field of plant biostimulants and considers the fundamental and innovative principles underlying this technology. The elucidation of the biological basis of biostimulant function is a prerequisite for the development of science-based biostimulant industry and sound regulations governing these compounds. The task of defining the biological basis of biostimulants as a class of compounds, however, is made more complex by the diverse sources of biostimulants present in the market, which include bacteria, fungi, seaweeds, higher plants, animals and humate-containing raw materials, and the wide diversity of industrial processes utilized in their preparation. To distinguish biostimulants from the existing legislative product categories we propose the following definition of a biostimulant as “a formulated product of biological origin that improves plant productivity as a consequence of the novel or emergent properties of the complex of constituents, and not as a sole consequence of the presence of known essential plant nutrients, plant growth regulators, or plant protective compounds.” The definition provided here is important as it emphasizes the principle that biological function can be positively modulated through application of molecules, or mixtures of molecules, for which an explicit mode of action has not been defined. Given the difficulty in determining a “mode of action” for a biostimulant, and recognizing the need for the market in biostimulants to attain legitimacy, we suggest that the focus of biostimulant research and validation should be upon proof of efficacy and safety and the determination of a broad mechanism of action, without a requirement for the determination of a specific mode of action. While there is a clear commercial imperative to rationalize biostimulants as a discrete class of products, there is also a compelling biological case for the science-based development of, and experimentation with biostimulants in the expectation that this may lead to the identification of novel biological molecules and phenomenon, pathways and processes, that would not have been discovered if the category of biostimulants did not exist, or was not considered legitimate.

The interest of biologists in boron (B) has largely been focused on its role in plants for which B was established as essential in 1923 (Warington, 1923[296]). Evidence that B has a biological role in other organisms was first indicated by the establishment of essentiality of B for diatoms (Smyth and Dugger, 1981[296]) and cyanobacteria (Bonilla et al., 1990[296]; Garcia‐Gonzalez et al., 1991[296]; Bonilla et al., 1997[296]). Recently, B was shown to stimulate growth in yeast (Bennett et al., 1999[296]) and to be essential for zebrafish (Danio rerio) (Eckhert and Rowe, 1999[296]; Rowe and Eckhert, 1999[296]) and possibly for trout (Oncorhynchus mykiss) (Eckhert, 1998[296]; Rowe et al., 1998[296]), frogs (Xenopus laevis) (Fort et al., 1998[296]) and mouse (Lanoue et al., 2000[296]). There is also preliminary evidence to suggest that B has at least a beneficial role in humans (Nielsen, 2000[296]). While research into the role of B in plants has been ongoing for 80 years it has only been in the past 5 years that the first function of B in plants has been defined. Boron is now known to be essential for cell wall structure and function, likely through its role as a stabilizer of the cell wall pectic network and subsequent regulation of cell wall pore size. A role for B in plant cell walls, however, is inadequate to explain all of the effects of B deficiency seen in plants. The suggestion that B plays a broader role in biology is supported by the discovery that B is essential for animals where a cellulose‐rich cell wall is not present. Careful consideration of the physical and chemical properties of B in biological systems, and of the experimental data from both plants and animals suggests that B plays a critical role in membrane structure and hence function. Verification of B association with membranes would represent an important advance in modern biology. For several decades there has been uncertainty as to the mechanisms of B uptake and transport within plants. This uncertainty has been driven by a lack of adequate methodology to measure membrane fluxes of B at physiologically relevant concentrations. Recent experimentation provides the first direct measurement of membrane permeability of B and illustrates that passive B permeation contributes sufficient B at adequate levels of B supply, but would be inadequate at conditions of marginal B supply. The hypothesis that an active, carrier mediated process is involved in B uptake at low B supply is supported by research demonstrating that B uptake can be stimulated by B deprivation, that uptake rates follow a Michaelis‐Menton kinetics, and can be inhibited by application of metabolic inhibitors. Since the mechanisms of element uptake are generally conserved between species, an understanding of the processes of B uptake is relevant to studies in both plants and animals. The study of B in plant biology has progressed markedly in the last decade and we are clearly on the cusp of additional, significant discoveries. Research in this field will be greatly stimulat...

Sedimentation velocity analytical ultracentrifugation is an important tool in the characterization of macromolecules and nanoparticles in solution. The sedimentation coefficient distribution c(s) of Lamm equation solutions is based on the approximation of a single, weight-average frictional coefficient of all particles, determined from the experimental data, which scales the diffusion coefficient to the sedimentation coefficient consistent with the traditional s approximately M(2/3) power law. It provides a high hydrodynamic resolution, where diffusional broadening of the sedimentation boundaries is deconvoluted from the sedimentation coefficient distribution. The approximation of a single weight-average frictional ratio is favored by several experimental factors, and usually gives good results for chemically not too dissimilar macromolecules, such as mixtures of folded proteins. In this communication, we examine an extension to a two-dimensional distribution of sedimentation coefficient and frictional ratio, c(s,f(r)), which is representative of a more general set of size-and-shape distributions, including mass-Stokes radius distributions, c(M,R(S)), and sedimentation coefficient-molar mass distributions c(s,M). We show that this can be used to determine average molar masses of macromolecules and characterize macromolecular distributions, without the approximation of any scaling relationship between hydrodynamic and thermodynamic parameters.

This article provides the context for a Special Issue of the Annals of Botany on 'Plant Nutrition for Sustainable Development and Global Health'. It provides an introduction to plant mineral nutrition and explains how mineral elements are taken up by roots and distributed within plants. It introduces the concept of the ionome (the elemental composition of a subcellular structure, cell, tissue or organism), and observes that the activities of key transport proteins determine species-specific, tissue and cellular ionomes. It then describes how current research is addressing the problems of mineral toxicities in agricultural soils to provide food security and the optimization of fertilizer applications for economic and environmental sustainability. It concludes with a perspective on how agriculture can produce edible crops that contribute sufficient mineral elements for adequate animal and human nutrition.

The protein refractive index increment, dn/dc, is an important parameter underlying the concentration determination and the biophysical characterization of proteins and protein complexes in many techniques. In this study, we examine the widely used assumption that most proteins have dn/dc values in a very narrow range, and reappraise the prediction of dn/dc of unmodified proteins based on their amino acid composition. Applying this approach in large scale to the entire set of known and predicted human proteins, we obtain, for the first time, to our knowledge, an estimate of the full distribution of protein dn/dc values. The distribution is close to Gaussian with a mean of 0.190 ml/g (for unmodified proteins at 589 nm) and a standard deviation of 0.003 ml/g. However, small proteins <10 kDa exhibit a larger spread, and almost 3000 proteins have values deviating by more than two standard deviations from the mean. Due to the widespread availability of protein sequences and the potential for outliers, the compositional prediction should be convenient and provide greater accuracy than an average consensus value for all proteins. We discuss how this approach should be particularly valuable for certain protein classes where a high dn/dc is coincidental to structural features, or may be functionally relevant such as in proteins of the eye.

Multisite interactions and the formation of ternary or higher-order protein complexes are ubiquitous features of protein interactions. Cooperativity between different ligands is a hallmark for information transfer, and is frequently critical for the biological function. We describe a new computational platform for the global analysis of isothermal titration calorimetry (ITC) data for the study of binary and ternary multisite interactions, implemented as part of the public domain multimethod analysis software SEDPHAT. The global analysis of titrations performed in different orientations was explored, and the potential for unraveling cooperativity parameters in multisite interactions was assessed in theory and experiment. To demonstrate the practical potential and limitations of global analyses of ITC titrations for the study of cooperative multiprotein interactions, we have examined the interactions of three proteins that are critical for signal transduction after T-cell activation, LAT, Grb2, and Sos1. We have shown previously that multivalent interactions between these three molecules promote the assembly of large multiprotein complexes important for T-cell receptor activation. By global analysis of the heats of binding observed in sets of ITC injections in different orientations, which allowed us to follow the formation of binary and ternary complexes, we observed negative and positive cooperativity that may be important to control the pathway of assembly and disassembly of adaptor protein particles.

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