MINI REVIEW

published: 27 August 2015

doi: 10.3389/fpls.2015.00671

Biostimulants in agriculture
Patrick Brown 1* and Sebastian Saa 2
1
Department of Plant Sciences, University of California, Davis, Davis, CA, USA, 2 Escuela de Agronomía, Pontificia
Universidad Católica de Valparaíso, Quillota, Chile

Keywords: biostimulants, mode of action, plant signaling, stress response, endophytic microorganisms, microbial
extracts

The past decades have witnessed tremendous growth in the use of biostimulants in agriculture
and it is estimated that biostimulants will grow to $2 billion in sales by 2018 (Calvo et al., 2014).
Recognizing the need to establish a legal framework for the marketing and regulation of these
products the European biostimulants industry council (EBIC, 2012) defined plant biostimulants
as “containing substance(s) and/or micro-organisms whose function when applied to plants or the
rhizosphere is to stimulate natural processes to enhance/benefit nutrient uptake, nutrient efficiency,
tolerance to abiotic stress, and crop quality.”
There is a clear need to improve our understanding of biostimulant function so that the efficacy
of these materials can be improved and the industrial processes can be optimized. Determining
the function of this class of products, however, has proven to be immensely difficult (Khan et al.,
2009; Carvalhais et al., 2013; Rose et al., 2014). This is in large part due to the diversity of sources
of these materials and the complexity of the resulting product, which in most cases will contain
Edited by: a significant number of poorly characterized molecules. Since biostimulants are derived from an
Ebrahim Hadavi, incredibly diverse set of biological and inorganic materials (Calvo et al., 2014) including microbial
Islamic Azad University, Iran fermentations of animal or plant feedstock, living microbial cultures, macro, and micro-alga,
Reviewed by: protein hydrolysate, humic, and fulvic substances, composts, manures, food, and industrial wastes
Victoria Fernandez, prepared using widely divergent industrial manufacturing processes, it is illogical to assume that
Technical University of Madrid, Spain there is a single mode of action.
Ioannis S. Minas,
The definition of biostimulants adopted by EBIC specifies that these materials should not
Aristotle University of Thessaloniki,
function by virtue of the presence of essential mineral elements, known plant hormones or
Greece
disease suppressive molecules. Accepting this definition, we hypothesize that biostimulants benefit
*Correspondence:
plant productivity by interacting with plant signaling processes thereby reducing negative plant
Patrick Brown,
Department of Plant Sciences,
response to stress. This hypothesis recognizes the wealth of recent research demonstrating that
University of California, Davis, MS#2, plant response to stress is regulated by signaling molecules that may be generated by the plant or
One Shields Avenue, Davis, its associated microbial populations (Marasco et al., 2012; Bakker et al., 2014; Vandenkoornhuyse
CA 95616, USA et al., 2015). Biostimulants may either directly interact with plant signaling cascades or act through
phbrown@ucdavis.edu stimulation of endophytic and non-endophytic bacteria, yeast, and fungi to produce molecules of
benefit to the plant (Figure 1). The benefit of the biostimulant is derived from the reduction in
Specialty section: assimilates that are diverted to non-productive stress response metabolism.
This article was submitted to
In this research topic the effects of biostimulants on plant productivity is examined in 10 research
Crop Science and Horticulture,
papers. Colla et al. (2014), soil-applied a plant-derived protein hydrolosate and demonstrated
a section of the journal
Frontiers in Plant Science improved growth and nitrogen assimilation in seedlings of pea, tomato, and corn. The use of
giberrellic acid (GA) deficient mutants and classic auxin response treatments suggests this material
Received: 12 June 2015
benefits plant growth by mimicking the actions of indole acetic acid (IAA) and GA.
Accepted: 13 August 2015
Published: 27 August 2015 Ertani et al. (2014) observed the effects of alfalfa hydrolosate (AH) and red grape extract (RG)
on nitrogen metabolism and growth of pepper plants (Capsicum chinensis). Significant, dose
Citation:
Brown P and Saa S (2015)
dependent changes were observed in a wide range of sugars, phenols, and quarternary nitrogen
Biostimulants in agriculture. containing molecules. In almond grown under high nutrient supply conditions biostimulants
Front. Plant Sci. 6:671. derived from either seaweed or microbial fermentation of cereal grains, had a marked positive
doi: 10.3389/fpls.2015.00671 effect on shoot growth and leaf area (Saa et al., 2015). Under conditions of low nutrient supply

Frontiers in Plant Science | www.frontiersin.org 1 August 2015 | Volume 6 | Article 671

Brown and Saa Biostimulants in agriculture

mapping techniques (µ-Xray Florescence) the movement of

Zn to the phloem following application of a combination of
biostimulant and zinc sulfate was elegantly demonstrated. This
research did not determine if the addition of the biostimulant
enhanced Zn uptake by increasing Zn movement through the leaf
surface and subsequent transport of Zn to the phloem, or if the
enhanced transport was a result of increased sink strength as was
observed when this same product was used in Almond (Saa et al.,
2015).
Vergnes et al. (2014) used foliar application of an essential
oil derived from Gaultheria procumbens and demonstrated
significant induced resistance on Arabidopsis leaves inoculated
with the fungal pathogen C. higginsianum. The authors
concluded that the essential oil from G. procumbens could be
a valuable natural source of methyl salicylic acid (MeSA) for
biocontrol applications. The application of salicylic acid (SA) has
been shown to have negative effects on plant productivity either
as a result of direct toxicity or changes in allocation of assimilates
to plant defense responses. This response was also observed by
Ghazijahani et al. (2014) who noted that the negative effects of
SA can be mitigated by co-application of citric acid.
FIGURE 1 | Non-lethal stress is experienced to varying degrees by all
crop plants resulting in a loss of productivity as assimilates are
Many biostimulants contain simple and complex
diverted to stress response metabolism (top figure). It is hypothesized carbohydrates that when applied to plant may alter metabolism
that biostimulants interacting with plant signaling processes reduce the extent by directly acting as a source of energy for endophytic and
of negative plant response to stress and increase the allocation of biomass to non-endophytic microbial populations or acting as signaling
the harvested yield component.
molecules. The complexity of the roles of carbohydrates in
plant immunity was reviewed by Trouvelot et al. (2014), who
suggested that carbohydrates activate defense reactions by
the benefit was less significant though there was a marked pathogen associated molecular patterns (PAMPs), microbe
increase in rubidium uptake (an analog for K uptake). A associated molecular patterns (MAMPS), and damage associated
differential response to the application of a nitrophenolate based molecular patterns (DAMPs). The authors highlight the main
biostimulant (Przybysz et al., 2014) was observed with significant classes of carbohydrates that are involved in plant immunity
and consistent growth and photosynthesis improvements under (beta-glucans, chitin, pectin) and discuss how the degree of
drought and heavy metal stress (platinum) and inconsistent polymerization and types of oligosaccharides affects biological
growth benefit under non-stressed growth conditions. activity. This review further suggests that carbohydrates in
Evidence that biostimulants may enhance macro nutrient biostimulants may act by beneficially manipulating plant
uptake has been reported previously (Calvo et al., 2014; Rose signaling cascades.
et al., 2014) and have been ascribed to an effect on sink activity The great diversity of plant response to biostimulants
or stimulation of nitrogen metabolism. Foliar application of highlights the challenges faced by researchers. Many plant
a biostimulant derived from microbial fermentation of cereal responses to biostimulants cannot be explained by our current
grains (Tian et al., 2015) greatly enhanced the movement of understanding of plant processes and while this represents a
foliar applied zinc in sunflower. Using high resolution elemental challenge, it also presents a great opportunity.

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of microbial and plant based biostimulants increases growth and potassium Conflict of Interest Statement: The authors declare that the research was
uptake in almond (Prunus dulcis Mill. D. A. Webb). Front. Plant Sci. 6:87. doi: conducted in the absence of any commercial or financial relationships that could
10.3389/fpls.2015.00087 be construed as a potential conflict of interest.
Tian, S., Lu, L., Xie, R., Zhang, M., Jernstedt, J. A., Hou, D., et al. (2015).
Supplemental macronutrients and microbial fermentation products improve Copyright © 2015 Brown and Saa. This is an open-access article distributed under the
the uptake and transport of foliar applied zinc in sunflower (Helianthus annuus terms of the Creative Commons Attribution License (CC BY). The use, distribution or
L.) plants. Studies utilizing micro X-ray florescence. Front. Plant Sci. 5:808. doi: reproduction in other forums is permitted, provided the original author(s) or licensor
10.3389/fpls.2014.00808 are credited and that the original publication in this journal is cited, in accordance
Trouvelot, S., Héloir, M. C., Poinssot, B., Gauthier, A., Paris, F., Guillier, with accepted academic practice. No use, distribution or reproduction is permitted
C., et al. (2014). Carbohydrates in plant immunity and plant protection: which does not comply with these terms.

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