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Hormonal Regulation of Plant Growth and Development

  • William M Gray

PLOS

Published: September 14, 2004

  • https://doi.org/10.1371/journal.pbio.0020311
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Citation: Gray WM (2004) Hormonal Regulation of Plant Growth and Development. PLoS Biol 2(9): e311. https://doi.org/10.1371/journal.pbio.0020311

Copyright: © 2004 William M. Gray. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Abbreviations: ABA, abscisic acid; ARF, auxin response factor; BR, brassinosteroid; GA, gibberellin; HK, histidine kinase; IAA, indole-3-acetic acid; JA, jasmonic acid; SCF, SKP1/Cullin/F-box protein

Phytohormones: What Are they?

Plant growth and development involves the integration of many environmental and endogenous signals that, together with the intrinsic genetic program, determine plant form. Fundamental to this process are several growth regulators collectively called the plant hormones or phytohormones. This group includes auxin, cytokinin, the gibberellins (GAs), abscisic acid (ABA), ethylene, the brassinosteroids (BRs), and jasmonic acid (JA), each of which acts at low concentrations to regulate many aspects of plant growth and development.

With the notable exception of the steroidal hormones of the BR group, plant hormones bear little resemblance to their animal counterparts ( Figure 1 ). Rather, they are relatively simple, small molecules such as ethylene gas and indole-3-acetic acid (IAA), the primary auxin in the majority of plant species. The concept of plant hormones originates from a classical experiment on phototropism, the bending of plants toward light, carried out by Charles Darwin and his son Francis in 1880. The Darwins were able to demonstrate that when oat seedlings were exposed to a lateral light source, a transported signal originating from the plant apex promoted differential cell elongation in the lower parts of the seedling that resulted in it bending toward the light source. This signal was subsequently shown to be IAA, the first known plant hormone.

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A partial list of the responses elicited by each hormone is provided below. Ethylene gas promotes fruit ripening, senescence, and responses to pathogens and abiotic stresses. IAA (an auxin) regulates cell division and expansion, vascular differentiation, lateral root development, and apical dominance. Cytokinins are adenine derivatives first identified by their ability to promote cytokinesis. JA is a volatile signal that modulates pollen development and responses to pathogen infection. The BRs regulate cell expansion and photomorphogenesis (light-regulated development). GAs are diterpenoid compounds that promote germination, stem elongation, and the induction of flowering. ABA promotes seed dormancy and is involved in several stress signaling pathways.

https://doi.org/10.1371/journal.pbio.0020311.g001

What Do They Do?

Virtually every aspect of plant growth and development is under hormonal control to some degree. A single hormone can regulate an amazingly diverse array of cellular and developmental processes, while at the same time multiple hormones often influence a single process. Well-studied examples include the promotion of fruit ripening by ethylene, regulation of the cell cycle by auxin and cytokinin, induction of seed germination and stem elongation by GA, and the maintenance of seed dormancy by ABA. Historically, the effects of each hormone have been defined largely by the application of exogenous hormone. More recently, the isolation of hormone biosynthetic and response mutants has provided powerful new tools for painting a clearer picture of the roles of the various phytohormones in plant growth and development.

How Do They Work?

Plant biologists have been fascinated by the regulatory capacity of phytohormones since the time of their discovery, and the notion that hormone levels or responses could be manipulated to improve desired plant traits has long been an area of intense interest. Perhaps the best-known example of this is the isolation of dwarf varieties of wheat and rice that led to the “green revolution” in the second half of the 20th century, which is credited with saving millions of people around the globe from starvation. These dwarf varieties have shorter stems than wild-type, making these plants less susceptible to damage by wind and rain. The molecular isolation of these “dwarfing genes” has revealed that they encode components of the GA biosynthesis and response pathways ( Peng et al. 1999 ; Sasaki et al. 2002 ).

To elucidate the molecular mechanisms underlying phytohormone action, several researchers have utilized the genetically facile model plant Arabidopsis thaliana to isolate mutations that confer altered response to applied hormone. Molecular and biochemical analysis of the gene products defined by these mutations, coupled with expression studies aimed at identifying the downstream target genes that mediate hormonal changes in growth and development, has begun to unlock some of the mysteries behind phytohormone action. While no hormone transduction pathway is completely understood, we now have a rudimentary understanding of many of the molecular events underlying hormone action. Several reviews covering the individual hormone pathways in greater detail have recently been published ( Turner et al. 2002 ; Gomi and Matsuoka 2003 ; Himmelbach et al. 2003 ; Kakimoto 2003 ; Dharmasiri and Estelle 2004 ; Guo and Ecker 2004 ; Wang and He 2004 ).

Common Themes

Regulation by proteolysis has emerged as a resounding theme in plant hormone signaling. The ubiquitin-mediated degradation of key regulatory proteins has been demonstrated, or is at least likely, for all of the phytohormone response pathways ( Smalle and Vierstra 2004 ). In the case of auxin, the response pathway is normally subject to repression by a large family of transcriptional regulators called the Aux/IAA proteins ( Figure 2 ). These proteins dimerize with members of the auxin response factor (ARF) family of transcription factors, thus preventing ARFs from activating auxin-responsive genes ( Tiwari et al. 2004 ). Upon an auxin stimulus, an SCF (SKP1/Cullin/F-box protein) ubiquitin ligase ( Deshaies 1999 ) containing the TIR1 F-box protein ubiquitinates the Aux/IAA proteins, marking them for degradation by the 26S proteasome thereby de-repressing the response pathway ( Gray et al. 2001 ). The hormone promotes the Aux/IAA–TIR1 interaction; however, the molecular mechanisms behind this regulation are unclear. Most yeast and animal SCF substrates must be post-translationally modified, usually by phosphorylation, before they are recognized by their cognate F-box protein. Despite numerous efforts to identify auxin-induced modification of Aux/IAA proteins, no such signal has been discovered, raising the distinct possibility that auxin uses a novel mechanism to regulate SCF–substrate interactions.

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(A) Wild-type Arabidopsis thaliana and the axr2-1 mutant. axr2-1 is a dominant gain-of-function mutation in an Aux/IAA gene that confers reduced auxin response. The mutant axr2-1 protein constitutively represses auxin response because it cannot be targeted for proteolysis by the SCF TIR1 ubiquitin ligase. The effect of the mutation on AXR2 stability is shown in a pulse-chase experiment (inset). Wild-type and axr2-1 seedlings were labeled with 35 S-methionine and AXR2/axr2-1 protein was immunoprecipitated either immediately after the labeling period (t = 0) or following a 15-minute chase with unlabeled methionine (t = 15).

(B) A simplified model for auxin response. In the absence of an auxin stimulus, Aux/ IAA proteins inhibit ARF transcriptional activity by forming heterodimers. Auxin perception (by an unknown receptor) targets the Aux/IAA proteins to the SCF TIR1 complex, resulting in their ubiquitination and degradation, thereby de-repressing the ARF transcription factors. Among the ARF targets are the Aux/IAA genes themselves, which produce nascent Aux/IAA proteins that restore repression upon the pathway in a negative feedback loop.

https://doi.org/10.1371/journal.pbio.0020311.g002

Ethylene and cytokinin are both perceived by receptors sharing similarity to bacterial two-component regulators. Common in prokaryotes, but apparently restricted to plants and fungi in eukaryotes, these modular signaling systems involve a membrane-bound receptor containing an intracellular histidine kinase (HK) domain ( Wolanin et al. 2002 ). Ligand binding activates the kinase, resulting in autophosphorylation and initiation of a series of phosphotransfer reactions that culminates with the activation of a response regulator protein that functions as the effector component of the pathway. Cytokinin signaling appears to largely follow this paradigm ( Kakimoto 2003 ). Ethylene response, however, appears more complex ( Guo and Ecker 2004 ).

Ethylene is perceived by a family of five receptors. ETR1 and ERS1 contain a consensus HK domain, however, the HK domains of ETR2, ERS2, and EIN4 are degenerate and lack elements necessary for catalytic activity. This fact, together with studies of “kinase-dead” mutants of ETR1 , suggests that HK activity is not required for ethylene response. Mutations that abolish ethylene binding in any of the five receptor genes are dominant and confer ethylene insensitivity, indicating that the receptors function as negative regulators of the ethylene pathway.

Genetic and molecular studies have positioned these receptors upstream of the Raf-like MAP kinase kinase kinase, CTR1, which interacts with the receptors and also acts as a negative regulator ( Figure 3 ). The integral membrane protein, EIN2, and the transcription factors EIN3 and EIL1 are positive regulators of ethylene signaling downstream of CTR1. Current models propose that hormone binding inactivates the receptors, thus resulting in down-regulation of CTR1 activity. Since the identification of CTR1, biologists have speculated that a MAP kinase cascade may be involved. Only recently, however, have putative MAP kinase kinase and MAP kinase components of the ethylene pathway been identified ( Chang 2003 ). Interestingly, these kinases appear to positively regulate ethylene response, suggesting that CTR1 must inhibit their function. If so, this would represent a novel twist on the traditional MAP kinase signaling paradigm. Precisely how the ethylene signal is transduced to the EIN3 and EIL1 transcription factors remains unclear. However, the recent finding that ethylene stabilizes these transcription factors, which are targeted for degradation by an SCF complex in the absence of ethylene, clearly indicates a role for the ubiquitin pathway ( Guo and Ecker 2003 ; Potuschak et al. 2003 ). One of the known targets for EIN3 is the ERF1 transcription factor, which activates several genes involved in a subset of ethylene responses.

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Ethylene is perceived by a family of two-component receptors containing a consensus (unshaded) or degenerate (shaded) HK domain (H). Three of the receptors also contain a C-terminal receiver domain (R). The receptors negatively regulate ethylene response together with CTR1 in a complex on the endoplasmic reticulum membrane. Perception results in reduced receptor and CTR1 activities and activation of a MAP kinase kinase, which transmits the signal through the EIN2 membrane protein, ultimately resulting in the activation of a transcriptional cascade in the nucleus. The EIN3 and EIL1 transcription factors regulate primary response genes including ERF1 , which activates a subset of secondary ethylene-induced genes involved in defense responses. EIN3/EIL1 abundance is regulated in an ethylene-dependent manner by SCF complexes containing F-box proteins encoded by the ethylene-induced genes EBF1 and EBF2 . Positive- and negative-acting components of the pathway are indicated in green and red, respectively. Solid lines indicate regulation that is likely to be through direct interactions. Dotted lines indicate speculative interactions based on genetic studies.

https://doi.org/10.1371/journal.pbio.0020311.g003

Signal Integration and Combinatorial Control

Long ago, plant physiologists noted the apparent antagonistic interactions between some of the phytohormones, such as between auxin and cytokinin in the regulation of root–shoot differentiation and between GA and ABA in germination. Other processes are synergistically regulated by multiple hormones. While it has long been obvious that hormones do not function in discrete pathways, but rather exhibit extensive cross-talk and signal integration with each other and with environmental and developmental signaling pathways, the molecular basis for such coordinated regulation has been unclear. Several recent findings have begun to elucidate the molecular details of some of these events.

One example of such signal integration was recently described for the ethylene and JA pathways ( Lorenzo et al. 2003 ). Genetic studies had previously implicated both hormones as important regulators of pathogen defense responses, as well as of the wounding response and other stress-related pathways. Additionally, microarray analysis has identified a large number of genes that are responsive to both hormones. The ERF1 transcription factor was recently found to be an intersection point for these two signaling pathways ( Lorenzo et al. 2003 ). Like ethylene, JA rapidly induces ERF1 expression, and treatment with both hormones synergistically activates ERF1 . Induction of ERF1 by both hormones alone or in combination is dependent upon both signaling pathways, and constitutive overexpression of ERF1 rescues the defense-response defects of both ethylene- and JA-insensitive mutants. These findings suggest that ERF1 represents one of the first signaling nodes identified in the complex web of hormonal cross-talk.

The auxin and BR pathways also appear to converge and mutually regulate some developmental processes. Both hormones promote cell expansion, and microarray studies have revealed that as many as 40% of all BR-induced genes are also up-regulated by auxin ( Goda et al. 2004 ; Nemhauser et al. 2004 ). BR is perceived by the cell surface receptor kinase BRI1 ( Wang and He 2004 ). The SHAGGY/GSK3-type kinase BIN2 acts as a negative regulator of the pathway downstream of the receptor. In the absence of a BR signal, BIN2 phosphorylates the transcription factors BES1 and BZR1, targeting them for proteolysis by the 26S proteasome. Upon a BR stimulus, BIN2 is inactivated, allowing BES1 and BZR1to accumulate in the nucleus, where they are presumably involved in regulating BR-responsive genes.

Using combined genetic, physiological, and genomic approaches, Nemhauser and colleagues (2004) were able to demonstrate that auxin and BR regulate Arabidopsis hypocotyl (embryonic stem) elongation in a synergistic and interdependent fashion. Elevating endogenous auxin levels rendered plants more sensitive to BR application in hypocotyl elongation assays, and this response was dependent upon both the auxin and BR signaling pathways. Genetic studies suggest that the convergence of these two pathways occurs at a late point in hormone signaling, perhaps at the promoters of the many genes responsive to both hormones. In support of this notion, bioinformatic analysis identified distinct sequence elements that were enriched specifically in the promoters of auxin-induced, BR-induced, and auxin/BR-induced genes.

Many Unanswered Questions

While great strides have been made in recent years in understanding the molecular basis of phytohormone action, many fundamental questions remain. Receptors and other upstream signaling components remain to be identified for the majority of the phytohormones. Equally important are the elucidation of hormonal networks and the integration of these networks with the morphogenetic program, such that our understanding of hormone action can be placed in a developmental context.

Acknowledgments

The author wishes to thank members of his lab for helpful comments on this manuscript. Work in the author's laboratory on auxin response is supported by National Institutes of Health grant GM067203 and the Mcknight Foundation.

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Plant hormones in growth and development

Plants use hormones to regulate virtually every aspect of their growth and development. Such signaling molecules have long attracted the attention of plant scientists and chemical modulation of hormone biology is widely used in agriculture. Despite the long history of plant hormone biology, the wide variety of signaling molecules plants use and the diversity of processes they control inevitably means that our understanding remains incomplete. Technological advances and ongoing interest in biotechnological applications mean that plant biologists continue to advance the state of art in hormone research. Therefore, the editors of Nature Communications ,  Communications Biology,  and Scientific Reports invite submissions that showcase new advances in understanding the metabolism, transport and signaling of plant hormones and how this impacts growth and development.

Potted Calla Lily (Zantedeschia sp.) on windowsill. Image two of two illustrating phototropism, the bending of plants toward a source of light. Auxins, chemicals which stimulate cell growth in plants, migrate to the less-lit part of the stem and cause cells to elongate, bending the plant toward the light source to maximize light exposure and, therefore, photosynthesis.

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Impacts of Plastics on Plant Development: Recent Advances and Future Research Directions

Enikő mészáros.

1 Department of Plant Biology, University of Szeged, Közép fasor 52, H6726 Szeged, Hungary

Attila Bodor

2 Department of Biotechnology, University of Szeged, Közép fasor 52, H6726 Szeged, Hungary; uh.degezs-u.oib@arodob (A.B.); uh.crb@ierep (K.P.)

3 Institute of Biophysics, Biological Research Centre, Temesvári krt. 62, H6726 Szeged, Hungary

Etelka Kovács

Sarolta papp, kamilla kovács, katalin perei, gábor feigl.

Plastics have inundated the world, with microplastics (MPs) being small particles, less than 5 mm in size, originating from various sources. They pervade ecosystems such as freshwater and marine environments, soils, and the atmosphere. MPs, due to their small size and strong adsorption capacity, pose a threat to plants by inhibiting seed germination, root elongation, and nutrient absorption. The accumulation of MPs induces oxidative stress, cytotoxicity, and genotoxicity in plants, which also impacts plant development, mineral nutrition, photosynthesis, toxic accumulation, and metabolite production in plant tissues. Furthermore, roots can absorb nanoplastics (NPs), which are then distributed to stems, leaves, and fruits. As MPs and NPs harm organisms and ecosystems, they raise concerns about physical damage and toxic effects on animals, and the potential impact on human health via food webs. Understanding the environmental fate and effects of MPs is essential, along with strategies to reduce their release and mitigate consequences. However, a full understanding of the effects of different plastics, whether traditional or biodegradable, on plant development is yet to be achieved. This review offers an up-to-date overview of the latest known effects of plastics on plants.

1. Introduction

The planet boundary concept defines the limits that humanity must not exceed in order to not endanger the favorable conditions in which it has been able to develop and live sustainably in a safe ecosystem [ 1 ]. In 2009, nine specific boundaries were established, including climate change, the loss of biosphere integrity, the disruption of nitrogen and phosphorus biogeochemical cycles, land use changes, ocean acidification, freshwater consumption, stratospheric ozone depletion, increasing aerosols in the atmosphere, and chemical pollution [ 2 ]. The last one encompasses the release of novel entities such as synthetic organic pollutants, heavy metal compounds, and plastic pollution into the environment [ 2 ].

After a temporary halt due to the COVID-19 pandemic in 2020, global plastic production rose to 390.7 million tons in 2022 [ 3 ], with projections indicating that its usage will reach 1231 million tons by 2060 [ 4 ]. Fossil-based plastics accounted for 90.2% of the world’s production in 2021, while bio-based/bio-attributed plastics and post-consumer recycled plastics comprised 8.3% and 1.5% of global plastic production, respectively [ 4 ]. In China, the total production of plastic products exceeded one billion tons by the end of 2019, establishing itself as the world’s largest producer and consumer of plastic [ 5 ]. In 2021, China was responsible for almost one third of the world’s plastic production (32%), followed by North America with 18%, then the rest of Asia with 17%, Europe with 15%, and Africa and the Middle East with 8% [ 4 ].

Plastics are artificially produced and polymerized from various monomers [ 6 ]. The most commonly found plastics in the environment are polyethylene (PE), polyvinyl chloride (PVC), polypropylene (PP), polyethylene terephthalate (PET), and polystyrene (PS) [ 7 ]. These materials have a wide range of applications, including, but not limited to, smartphones, food packaging, and 3D printing, which stems from its design adaptability, affordability, ability to be shaped, lightweight nature, and biologically inert properties [ 8 ]. Consequently, plastic has the advantage of replacing traditional materials, such as paper, wood, or metal.

Once released into the environment, plastics pose a significant threat, due to their slow decomposition, which can take hundreds of years. The primary sources of plastic pollution are the fragmentation of larger plastic items, such as bags, bottles, and packaging materials. Over time, these items break down into smaller pieces, eventually becoming macroplastics (>2.5 cm), microplastics (<5 mm, MPs), or even nanoplastics (<100 nm or <1000 nm) [ 9 , 10 ]. MPs have two main sources: primary and secondary. The primary sources include drugs, paints, cosmetics, biomedical equipment, and other items, while the secondary sources refer to the mechanical, thermal, and biological degradation of macroplastics [ 11 ]. Other sources of MP pollution include the use of synthetic textiles, such as polyester and nylon, which shed microfibers during washing, and the use of cosmetics or personal care products that contain microbeads. Additionally, MPs can be released into the environment through the disposal of electronic waste, such as mobile phones and computers [ 12 ]. MPs can interact with other chemicals, leading to the accumulation of organic and inorganic pollutants on their surfaces [ 13 ]. Polychlorinated biphenyls (PCBs), polycyclic aromatic hydrocarbons (PAHs), antibiotics, and other chemicals pose a significant concern when they attach to MPs [ 13 ]. Consequently, MPs are categorized as emerging persistent pollutants that occur widely in various ecosystems.

Most plastic becomes waste shortly after use, and, therefore, a significant amount of plastic is constantly released into the environment. According to estimations, about 12.7 million tons of plastic has ended up in the marine environment so far [ 14 ], and MPs have been detected in the air [ 15 ] and in animals, where they can accumulate in various tissues, posing a long-term threat. Moreover, MPs can be transferred through the trophic food chain, reaching final consumers, including predators and humans [ 16 ]. Importantly, recent research has revealed the presence of MPs in human blood, with a measured total amount of 1.6 µg/L. The main compounds found were PET, PE, and PS. These results show that these particles are bioavailable and can be absorbed into the bloodstream of humans [ 17 ].

Considering the persistence and widespread distribution of MPs in the soil, it is necessary to recognize their potential impacts on terrestrial plants. Based on estimations, approximately 32% of all plastic produced ends up in the soil [ 18 ]; therefore, soils can be a much larger sink for plastics than salt water and freshwater [ 19 ]. Consequently, the presence of plastics in soil ecosystems will have an impact on the organisms that live there, including plants.

Exploring the relationship between plants and plastics is currently becoming a hot research topic; however, despite the growing body of published results, it is still challenging to draw definitive conclusions. After reviewing the available literature, it seems to be evident that the impacts of MPs on higher plants depend on various factors, such as the properties of the MPs, the specific plant species involved, and the surrounding environmental conditions. Under certain experimental conditions, MPs have induced no effects, or even positive effects, on higher plants. However, a larger body of evidence demonstrates that MPs can directly and indirectly impede the growth of higher plants. As a recently published paper [ 20 ] has already compiled the known effects of plastics on plants, this article aims to summarize the most recent experimental data, with a primary focus on the effects of plastics on the rhizosphere, particularly on plants.

2. Impact of Plastics on Soil, Plants, and Ecosystems

Soil, as the uppermost loose, fertile layer of the Earth’s surface, is essential for sustaining life. It provides plants (and thus animals and humans) with nutrients and water, while also storing and transforming materials [ 21 ]. To safeguard the healthy status of soil, models show that the concentration of MPs should not exceed the range of 2128–14,435 mg for each kg soil in order to keep up with half of the currently present soil biota or soil properties [ 19 ]. However, agricultural soil serves as a larger sink for MPs compared to water [ 22 ]. MPs can enter soils through various pathways, including sewage and sludge irrigation and residual mulching film decomposition [ 23 , 24 ]. An increased amount of MPs in the soil can directly alter the soil physicochemical properties, leading to reduced soil aeration, increased erosion, altered soil pH, and reduced nutrient availability for plants, and, hence, lower crop yields [ 25 , 26 ]. Additionally, the presence of MPs may alter the composition of soil microbial communities, reducing the diversity and abundance of beneficial microorganisms, which play essential roles in soil fertility and plant growth. Furthermore, MPs can act as carriers of other chemical contaminants in the soil, causing damage to plant and human health, when they reach the food chain [ 13 , 18 , 19 ].

Once in the soil, some MPs can be transported by external factors and enter surface water and groundwater via horizontal and vertical migration, while others are absorbed or accumulated by plants and soil-dwelling organisms [ 27 ]. In addition to the adverse effects of MPs on soil properties [ 12 ], several studies have also confirmed the extensive negative impact of MP accumulation on soil biology [ 28 ]. For example, MPs can disrupt the functional and structural diversity of soil microbial communities [ 29 ] and harm soil-dwelling animals, plants, and microorganisms by damaging DNA, inducing oxidative stress, and impairing metabolic processes [ 30 , 31 ]. However, these effects are often contradictory, and vary depending on factors such as MP shape and size, polymer type, degradability, or the presence of additives and impurities [ 28 ].

When it comes to MP contamination in soil, invertebrates, particularly earthworms, have received significant attention in studies, as they are vital components of the soil food chain and can be used to assess the toxicity of contaminants such as MPs in soil [ 19 ]. Earthworms have the ability to ingest MPs and reduce their size, easing their decomposition [ 32 ]. Reduced growth rates [ 33 ], immunological stress responses [ 34 ], damaged intestinal cells and DNA [ 35 ], and increased mortality [ 36 ] are among the negative outcomes caused by exposure to MPs. In a recently published study investigating on the potential harm of low-density polyethylene (LDPE) to earthworms ( Eisenia fetida ), Mondal et al. [ 37 ] found that, during the 28-day exposure period, no mortality or weight loss relative to the controls was observed. However, they noticed damage to the surfaces of the earthworms’ skin caused by exposure to the LDPE MPs. The induced skin surface damage and MP uptake can adversely affect the growth and reproduction of E. fetida after long-term MP exposure.

Under the influence of environmental factors, MPs can be broken down into nanoplastics (NPs), which are plastic particles smaller than a micron in size [ 38 ]. Similar to MPs, NPs also come from a variety of sources, including the decomposition of plastic products in the environment and the influx of microbeads and raw materials used in industry [ 39 ]. Significant amounts of NPs are present in our daily lives, derived, for example, from tire wear, washing textiles, and using personal hygiene products [ 40 , 41 , 42 ]. The increasing abundance of NPs poses potential environmental hazards, adding to the global problem of environmental pollution. Living organisms can absorb NPs from the environment, triggering various stress responses. Since plants play an important role in ecosystems, the bioaccumulation of NPs can be an entry point for plastics into the food chain [ 43 ]; therefore, it is of utmost importance to study and explore the interactions and mechanisms between NPs and plant [ 38 ]. NPs can enter plants through various pathways, such as root or foliar uptake. The root uptake pathway can be significant for terrestrial plants, such as crops grown in agricultural fields, since they rely on soil water for their survival [ 10 ]. Foliar uptake occurs when plastic particles are present in the air, and the plant leaves absorb them through atmospheric deposition. This pathway can be significant for plants growing in urban environments or near industrial areas with elevated air pollution [ 44 ].

Plants are essential for our ecosystem, providing food, oxygen, and numerous other benefits, which can be jeopardized by anthropogenic stressors. In recent years, the issue of plastic pollution has emerged as a growing concern [ 45 , 46 ]. While plastic waste is known to harm marine life [ 47 ], scientists have only recently begun to realize the impact of MP pollution on terrestrial ecosystems [ 48 ] and its effect on plants [ 49 ]. The potential impacts of MPs and NPs on plants and the issue of food safety concerns for crops have been discussed recently [ 49 ], and subsequent studies have confirmed that NPs can be absorbed by plant roots and translocated to the aboveground aerial tissues [ 50 ]. These studies have demonstrated that MPs and NPs can induce physiological changes in plants, such as a reduction in growth, photosynthesis, and antioxidant activity, as well as alterations in gene expression and root exudate profiles [ 49 , 51 ].

Currently, most research focuses on the distribution and potential toxicity of MPs/NPs in aquatic ecosystems and their transmission to human food sources. However, there is also an urgent need to investigate the potential harmful effects of MPs/NPs on terrestrial plants and crop production. Although some recent studies, such as those by Zhou et al. [ 52 ], Yin et al. [ 53 ], Okeke et al. [ 54 ], and Yadav et al. [ 55 ], have explored the impacts of MPs on agroecosystems, vascular plants, food chains, and plant growth, these studies offer only a superficial analysis, and there is a lack of explanation regarding the underlying causes (such as the antagonistic effect of MPs/NPs on plants) that contribute to the dysfunction of the food chain. Therefore, it is crucial to conduct an in-depth analysis of the potential issues related to the entry of plastic particles into plants, the subsequent weakening of plant defense mechanisms, the putative factors determining the toxicity of MPs/NPs, and their interference with food quality and quantity. Moreover, as research in this area is still in its early stages, there is limited information on how to mitigate the adverse effects of MPs/NPs in plants.

3. Effects of Plastics on Plant Development

Studies have shown that plastics generally have a negative effect on plant development, which might manifest in alterations in both germination and root or shoot growth. These changes, however, depend on several factors, including the environmental conditions, plant species, and plastic concentration. Several types of plastics have been tested, including PS, PE, PVC, and biodegradable plastics, which are summarized below.

3.1. Effects of Polystyrene on Plant Development

PS is a widely used synthetic polymer derived from aromatic hydrocarbons called styrene monomers. With an annual production volume of several million tons, it ranks among the most commonly utilized plastics. Although PS is naturally transparent, it can be dyed for various applications. These applications include protective packaging containers, lids, bottles, trays, cups, and disposable cutlery [ 56 ].

The relationship between PS and plants has been studied in a range of applied concentrations and experimental systems, the vast majority of which have demonstrated growth inhibition. The studies dealing with the effects of polystyrene mentioned in the text are summarized in Table 1 .

In realistic field conditions, as stressors are seldom found in isolation within the environment, MPs and NPs can coexist with arsenic (As) in the soil, potentially causing toxic effects on plant growth and escalating the accumulation of arsenic in plants throughout the food chain. Two studies centered on the impact of PS plastic fragments on rice ( Oryza sativa L.) plants, along with the absorption of As. The studies aimed to explore how MPs influence the overall As uptake in rice seedlings and the subsequent accumulation of As within the rice tissues. The growth responses were found to be dependent on the size of the NP/MP particles. Interestingly, the exposure of rice seedlings to MPs slightly mitigated the adverse effects of As on plant leaf growth and reduced root activity when compared to rice seedlings that were exposed to arsenic alone [ 57 ]. An additional study conducted by Xu et al. [ 58 ] revealed a noteworthy decline ( p < 0.001) in aboveground tissue biomass due to As treatment, as compared to the control group. This effect was more pronounced when both MP and As stresses were combined. However, the root biomass exhibited only minor alterations. These results indicate that the combined effect of MPs and As on plants differs from the individual effects of MPs or As alone. Furthermore, other studies reported root and shoot inhibition in the presence of PS MPs on rice [ 59 ], which is consistent with earlier findings [ 60 ].

Lettuce (Lactuca sativa L.) is a major food source grown worldwide. Wang et al. [ 61 ] observed the effects of PS plastic on lettuce growth and found that PS MPs can disturb the antioxidant system, change the gene expression in roots, and influence the root exudate profiles. MP stress increased the expression of genes involved in different antioxidant systems at different times in the roots and leaves (ascorbic acid, terpenoids, flavonoids, and sphingolipids). The solutions with higher MP concentrations further inhibited lettuce growth compared to the controls, and the fresh leaf weight, plant height, and number of leaves were significantly reduced in all of the plants grown in the presence of MPs ( p < 0.05) [ 61 ].

The response of tomato ( Lycopersicon esculentum L.) plants to PS was investigated in plants grown in a hydroponic medium. Shi et al. [ 62 ] conducted an experiment with 13 treatment groups, including PS concentrations ranging from 0.1 to 1 mg/L, with tomato seedlings for 14 days. The results showed that PS MPs inhibit tomato plant growth and cause severe oxidative stress. Several treatments reduced the length of the shoot and root of the tomatoes and also affected some important metabolic pathways, including the tricarboxylic acid cycle and glutathione metabolism [ 62 ].

There are also some studies with mung bean ( Vigna radiata L.) and onion ( Allium cepa L.) plants in pot conditions. These studies have demonstrated that PS had a more negative effect on root growth in mung beans compared to onions. Biba et al. [ 63 ] reported that there were no significant changes in onion root growth after exposure to any of the tested PS MP concentrations, compared to the control, although the highest concentration (1 g/L) caused a slight decrease in length. On the other hand, Chen et al. [ 64 ] found that any applied concentration of PS MPs significantly reduced the growth characteristics of mung bean plants, including the shoot and root growth. PS also had a negative effect on the root and shoot growth of water spinach ( Ipomoea aquatica Forssk.) and dandelion ( Taraxacum asiaticum Dahlst) plants in hydroponic conditions but had no significant effect on their seed germination [ 65 , 66 ].

On the other hand, PS did not significantly reduce any root or shoot growth parameters in corn ( Zea mays L.) plants [ 67 ], and no negative effects were detected in soybean ( Glycine max L.) plants either [ 68 ]. Conversely, when watermilfoil ( Myriophyllum verticillatum L.) was tested, PS MPs had a negative effect on shoot growth [ 69 ]. Interestingly, a recently conducted study showed that the root and shoot growth of corn plants were inhibited by PS [ 70 ]. Another experiment showed that PS had a negative effect on root growth in soybean plants, while no changes were observed in shoot growth [ 71 ]. However, a recent study found a slight effect on seed germination and sprout growth in soybean plants [ 68 ].

The effect of polystyrene-based plastics on the development of plants. “−” represents inhibition, “0” marks no change, while “n.d.” marks parameters not determined.

3.2. Effects of Polyethylene Plastics on Plant Development

PE is the most widely used plastic, mainly by the packaging industry. It has several types, such as high-density polyethylene (HDPE), medium-density polyethylene (MDPE), low-density polyethylene (LDPE), and cross-linked polyethylene (XLPE/PEX) [ 72 ].

The impact of PE on plant development has been explored in several recent studies under different conditions ( Table 2 ). Numerous studies have shown that PE negatively affects both the root and shoot growth of corn plants [ 73 , 74 , 75 ]. Additionally, a significant reduction in root and shoot growth has been observed in cucumber ( Cucumis sativus L.) and water moss ( Salvinia auriculata Aubl.) plants in hydroponic experiments [ 76 , 77 ]. An experiment with lentil ( Lens culinaris Medik) plants demonstrated that their germination was inhibited by PE [ 78 ]. Moreover, a study found that PE significantly increased the shoot growth of lettuce, barley ( Hordeum vulgare L.), cucumber, and tomato ( Solanum lycopersicum L.) plants [ 50 , 79 ]. Conversely, according to another study, root development was reduced by PE in cucumber and tomato plants [ 79 ].

The effect of polyethylene-based plastics on the development of plants. “+” indicates growth induction, “0” means no change, “−” represents inhibition, while “n.d.” marks parameters not determined.

The effect of HDPE on plants has also been researched. For instance, barley, sand couch-grass ( Thinopyrum junceum L.), and ice plant ( Carpobrotus sp.) plants are sensitive to HDPE MPs stress, as it inhibits their root and shoot growth [ 80 , 81 ].

3.3. Effects of Polyvinyl Chloride Plastics on Plant Development

PVC is a chemically inert material that can exist in both flexible and rigid forms. Rigid PVC is easily machinable, thermoformable, weldable, and can even be bonded using solvents. PVC can also be machined using standard metalworking tools and is less difficult to machine to tight tolerances and finishes. PVC resins are commonly blended with other additives, such as impact modifiers and stabilizers, to create a wide range of PVC-based materials [ 82 ].

Several recent studies have demonstrated the negative effects of PVC MPs on plant development in different growing media, which are presented in Table 3 . PVC has been found to inhibit the growth of corn and plumed cockscomb ( Celosia argentea L.) roots and shoots [ 83 , 84 ]. However, there was no significant effect observed in sweet potato ( Ipomoea batatas L.) plants [ 85 ]. In duckweed ( Spirodela polyrhiza L.) plants, PVC MPs were found to have a greater impact on root extension than shoot growth [ 86 , 87 ]. Moreover, Song et al. [ 88 ] examined the effects of PVC plastic fragments on rice and found a negative effect on root growth but no significant impact on shoot development.

The effect of polyvinyl-chloride-based plastics on the development of plants. “0” means no change, “−” represents inhibition, while “n.d.” marks parameters not determined.

3.4. Effects of Biodegradable Plastics on Plant Development

Bioplastics are materials or products derived from biomass that are driving the evolution of plastics. These bio-based plastic products are typically derived from sources such as corn, sugarcane, or cellulose. They offer two main advantages over conventional products. Firstly, using biomass as a raw material saves fossil resources, as it is renewable and has the unique potential to be CO 2 neutral. Additionally, certain types of bioplastics exhibit biodegradability as an additional property [ 89 ].

The effect of bio-based plastics on plant development has been examined in several studies recently ( Table 4 ). Corn appears to be a particularly sensitive species to biodegradable plastics such as polybutylene adipate terephthalate (PBAT) and polylactic acid (PLA), as indicated by reduced root and shoot growth [ 73 , 74 ]. Poly(3-hydroxybutyrate-co-3-hydroxyvalerate) PHBV, a microbial biopolymer with excellent biocompatibility and biodegradability, has been shown to negatively impact corn root and shoot growth in soil [ 90 ]. Furthermore, pot experiments with basil ( Ocimum basilicum L.), sand couch-grass, and ice plant plants have demonstrated that biodegradable plastics such as corn-starch-based bioplastic and starch-based polymers inhibited root and shoot growth [ 81 , 91 ]. In the case of sorghum ( Sorghum saccharatum L.), garden cress ( Lepidium sativum L.), and white mustard ( Sinapis alba L.) plants, Liwarska-Bizukojc et al. [ 92 ] studied the effect of bio-based plastics such as PLA and polyhydroxybutyrate (PHB). The results indicated a significant reduction in both root and shoot growth. However, there were no statistically significant differences in germination efficiency between tests conducted with and without plastic particles in the soil. Interestingly, in red cherry tomato ( Lycopersicon esculentum Mill.) and lettuce plants, plastic particles such as PBAT and PLA inhibited germination and shoot growth as well [ 93 ]. Additionally, PLA inhibited the root and shoot development of cucumber and barley plants [ 76 , 80 ].

The effect of bio-based plastics on the development of plants. “0” indicates no change, “−” represents inhibition, while “n.d.” marks parameters not determined.

4. Conclusions

In summary, the effects of plastics on plant development are increasingly being investigated, and it is evident that different types of plastic can have varying impacts on plants. Consistent with previous reviews [ 20 , 59 , 94 , 95 ], the plant responses to plastics remain predominantly uniform. Most recent studies demonstrate growth inhibition, while only a handful of species exhibit a positive growth response, particularly in the presence of PE ( Figure 1 ). These additions include barley [ 79 ], cucumber, tomato, and lettuce plants [ 50 ], adding to previously identified species such as Allium fistulosum , T. aestivum , L. sativum , Calamagrostis epigejos , and Hieracium pilosella (reviewed in [ 20 ]).

An external file that holds a picture, illustration, etc.
Object name is plants-12-03282-g001.jpg

Plant growth responses to polystyrene, polyethylene, polyvinyl-chloride, and biodegradable plastics.

However, the aggregated results indicate that plastics, including PS, PE, PVC, and biodegradable variants, have predominantly detrimental effects on plant growth, particularly in relation to root and shoot development.

PS has consistently emerged as an inhibitor of plant growth, as evidenced by studies revealing diminished root and shoot expansion across various plant species. Likewise, PE exhibits inhibitory traits affecting crops such as maize, cucumber, and water moss, leading to compromised root and shoot proliferation. PVC’s negative impact on plant development is also evident, as demonstrated by reduced root growth in rice and plumed cockscomb plants. Despite their touted environmental friendliness, biodegradable plastics also manifest deleterious consequences for plant growth. Corn, basil, sand couch-grass, and ice plant display suppressed root and shoot development in the presence of such plastics. Moreover, PBAT and PLA hinder the germination and shoot growth of red cherry tomatoes and lettuce plants.

As previously summarized [ 20 , 59 , 94 , 95 ], the mechanisms through which microplastics affect the performance of higher plants are both diverse and intricate ( Figure 2 ). One of the predominant outcomes triggered by MPs in plants is oxidative stress, as evidenced by the increased generation of reactive oxygen species and heightened activity of antioxidant enzymes (as reviewed by [ 59 , 94 , 95 ]). Similarly, a common feature of the results presented in the reviewed papers is the disruption of oxidative homeostasis. This alteration implies its potential involvement in shaping the plant growth responses to plastics, similar to what has been documented for numerous other abiotic stressors. Furthermore, nutrient uptake and metabolism have often been cited as plausible explanations for growth reduction (see [ 59 , 94 , 95 ]), and the most recent evidence collected in this paper also support this possibility. In addition, the effect of plastic on photosynthetic activity is well-documented [as reviewed in [ 59 , 95 ]], a fact confirmed by several of the recent studies collected in this paper.

An external file that holds a picture, illustration, etc.
Object name is plants-12-03282-g002.jpg

Mechanisms through which microplastics affect the growth and development of plants.

However, it is crucial to recognize that the precise processes, triggers, and outcomes of these phenomena remain unclear, highlighting the need for further research. It is also important to note that the effects of plastics on plant development may vary depending on factors such as plastic type, concentration, plant species, and environmental conditions. Nevertheless, the overall findings highlight the potential harm that plastic pollution, both conventional and biodegradable, can have on plant growth and ecosystem health.

Further research is needed in order to better understand the mechanisms underlying the negative effects of plastics on plants and to develop mitigation strategies. Most importantly, efforts should be made to reduce plastic pollution and promote the use of sustainable alternatives in order to ensure the health and sustainability of our ecosystems and ensure the continued provision of essential ecosystem services by plants. Overall, addressing plastic pollution and its impact on plant development is critical for the conservation of biodiversity, food security, and the overall well-being of our planet.

5. Future Directions

Future research directions in this field involve the study of the following aspects:

  • Interactions with different stressors: The interactions between plastics and other stressors, such as heavy metals or chemicals (which can be adsorbed in MPs), need to be explored further. Understanding how plastics interact with other environmental factors can provide insights into their combined effects on plant growth and development.
  • Mechanisms of action: Studying the mechanisms through which plastics exert their negative effects on plants is crucial. This includes investigating how plastics are taken up by plants, their impact on cellular processes, and the disruption of plant physiology. Elucidating these mechanisms will help in designing targeted mitigation strategies.
  • Species-specific responses: Different plant species may exhibit varying sensitivities to plastics. Further research should focus on a wide range of plant species in order to better understand the species-specific responses to different types of plastics and concentrations.
  • Long-term effects: Most of the studies conducted so far have focused on short-term effects. It is important to investigate the long-term consequences of plastic exposure on plant growth, reproduction, and overall ecosystem health. Long-term studies can provide valuable insights into the persistence and cumulative effects of plastics on plants.
  • Field studies: While many studies have been conducted under controlled laboratory conditions, field studies are necessary to assess the real-world impacts of plastics on plant development. Field experiments can consider the complex interactions of plants with their natural environment, including soil composition, nutrient availability, and microbial communities.
  • Biodegradable plastics: Further research is needed to evaluate the environmental fate and potential ecological impacts of biodegradable plastics. Understanding their decomposition rates, byproducts, and effects on plant growth will help us to determine their suitability as alternatives to conventional plastics.

Acknowledgments

The authors would like to express their gratitude toward Erika Pál for the excellent administrative support.

Funding Statement

This work was supported by the National Research, Development and Innovation Office (NKFIH FK 142475) and by the János Bolyai Research Scholarship of the Hungarian Academy of Sciences (Grant no. BO/00181/21/4). G.F. was supported by the New National Excellence Program of the Ministry of Human Capacities (UNKP-22-5-SZTE-532).

Author Contributions

Conceptualization, A.B., E.K. and G.F.; investigation, E.M., A.B., E.K., S.P., K.K., K.P. and G.F.; writing—original draft preparation, E.M.; writing—review and editing, A.B., E.K., S.P., K.K., K.P. and G.F.; supervision, G.F.; funding acquisition, G.F. All authors have read and agreed to the published version of the manuscript.

Conflicts of Interest

The authors declare no conflict of interest.

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Dynamic DNA Methylation in Plant Growth and Development

Affiliations.

  • 1 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 2 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 3 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 4 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 5 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 6 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 7 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 8 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • 9 Department of Plant and Microbial Biology, North Carolina State University, Raleigh, NC 27695, USA. [email protected].
  • 10 Plants for Human Health Institute, North Carolina State University, North Carolina Research Campus, Kannapolis, NC 28081, USA. [email protected].
  • 11 Plant Genetics Research Unit, Donald Danforth Plant Science Center, Midwest Area, Agricultural Research Service, US Department of Agriculture, St. Louis, MO 63132, USA. [email protected].
  • 12 Department of Biology, Saint Louis University, St. Louis, MO 63103, USA. [email protected].
  • PMID: 30041459
  • PMCID: PMC6073778
  • DOI: 10.3390/ijms19072144

DNA methylation is an epigenetic modification required for transposable element (TE) silencing, genome stability, and genomic imprinting. Although DNA methylation has been intensively studied, the dynamic nature of methylation among different species has just begun to be understood. Here we summarize the recent progress in research on the wide variation of DNA methylation in different plants, organs, tissues, and cells; dynamic changes of methylation are also reported during plant growth and development as well as changes in response to environmental stresses. Overall DNA methylation is quite diverse among species, and it occurs in CG, CHG, and CHH (H = A, C, or T) contexts of genes and TEs in angiosperms. Moderately expressed genes are most likely methylated in gene bodies. Methylation levels decrease significantly just upstream of the transcription start site and around transcription termination sites; its levels in the promoter are inversely correlated with the expression of some genes in plants. Methylation can be altered by different environmental stimuli such as pathogens and abiotic stresses. It is likely that methylation existed in the common eukaryotic ancestor before fungi, plants and animals diverged during evolution. In summary, DNA methylation patterns in angiosperms are complex, dynamic, and an integral part of genome diversity after millions of years of evolution.

Keywords: DNA methylation; development; dynamics; epigenetics; gene expression; methylome; plant; seed; transposable element.

Publication types

  • DNA Methylation*
  • Epigenesis, Genetic
  • Gene Expression Regulation, Developmental
  • Gene Expression Regulation, Plant
  • Plant Development / genetics*

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Study brings scientists a step closer to successfully growing plants in space

New, highly stretchable sensors can monitor and transmit plant growth information without human intervention, report University of Illinois Urbana-Champaign researchers in the journal Device.

The polymer sensors are resilient to humidity and temperature, can stretch over 400 percent while remaining attached to a plant as it grows and send a wireless signal to a remote monitoring location, said  chemical and biomolecular engineering  professor Ying Diao, who led the study with  plant biology  professor and department head Andrew Leakey.

Ying standing in lab and Andrew sitting in office

The study details some of the early results of a NASA grant  awarded to Diao  to investigate how wearable printed electronics will be used to make farming possible in space.

“This work is motivated by the needs of astronauts to grow vegetables sustainably while they are on long missions,” she said.

Diao’s team approached this project using an Earth-based laboratory to create a highly dependable, stretchable electronic device – and its development did not come easily, she said.

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Surfactin as a multifaceted biometabolite for sustainable plant defense: a review

  • Published: 26 April 2024

Cite this article

research on plant growth and development

  • Mohadeseh Hassanisaadi   ORCID: orcid.org/0000-0002-5626-6583 1  

Plants face numerous challenges in their ongoing battle against pests and diseases. Traditional protection methods often involve synthetic pesticides, which have a detrimental impact on the environment and human health. However, the quest for eco-friendly and sustainable solutions has brought surfactin into the spotlight as a promising defender of plants. Surfactin, a biometabolite produced by Bacillus spp., has gained attention due to its multifaceted properties contributing to plant defense. This review highlights the eco-friendly nature of surfactin and explores its notable functions as an antimicrobial agent, the ability to modulate plant defense mechanisms, enhance colonization and biofilm formation of antagonists, and ultimately promote plant growth. Furthermore, the environmentally friendly characteristics of surfactin, such as its biodegradability and low toxicity, make it an ideal candidate for sustainable plant protection strategies. The potential applications and challenges in utilizing surfactin as an eco-friendly defender of plants are discussed, providing insights for future research and the development of innovative and sustainable agricultural practices.

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Hassanisaadi, M. Surfactin as a multifaceted biometabolite for sustainable plant defense: a review. J Plant Pathol (2024). https://doi.org/10.1007/s42161-024-01645-9

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    Plant growth and development is the process by which plants respond to environmental signals and stimuli, such as light, temperature, and nutrients, to regulate their growth and development. ... International Journal of Aquaculture Research and Development . DOI: 10.14302/issn.2691-6622.ijar-20-3312. Full-text HTML . Download as PDF. Download ...

  23. Role of Nitrogen for Plant Growth and Development: A review

    Abstract and Figures. All plants utilize nitrogen (N) in the form of NO3- and NH4+. It is most imperative element for proper growth and development of plants which significantly increases and ...

  24. Surfactin as a multifaceted biometabolite for sustainable plant defense

    Effects of surfactin on plant growth and development. ... Furthermore, surfactin stimulates root development. Research has shown that surfactin stimulates root growth and development by increasing the length and branching of roots, resulting in a larger root system. Plants with a larger root system may absorb a greater amount of water and ...

  25. The Effect of Humic Substances on the Colony Growth and Conidial ...

    The development of sustainable agricultural production involves the use of new plant protection products, with low toxicity to non-target organisms and high biodegradability. The aim of this study was to investigate the effect of commercially available preparations containing humic substances, in comparison with pure humic acids, on the growth and germination of spores of entomopathogenic ...

  26. Serine proteinase inhibitors from

    Research Article. Serine proteinase inhibitors from Nicotiana benthamiana, a non-preferred host plant, inhibit the growth of Myzus persicae (green peach aphid) Honglin Feng, Corresponding Author. ... requiring the urgent development of novel strategies. Plant proteinase inhibitors (PINs) are small proteins that protect plants against pathogens ...