Heavy metals can represent a threat to the health of aquatic ecosystems. Unlike organic chemicals, heavy metals cannot be eliminated by natural processes such as their degradation into less toxic compounds, and this creates unique challenges for their remediation from soil, water, and air. Phytoremediation, defined as the use of plants for the removal of environmental contaminants, has many benefits compared to other pollution-reducing methods. Phytoremediation is simple, efficient, cost-effective, and environmentally friendly because it can be carried out at the polluted site, which simplifies logistics and minimizes exposure to humans and wildlife. Macrophytes represent a unique tool to remediate diverse environmental media because they can accumulate heavy metals from contaminated sediment via roots, from water via submerged leaves, and from air via emergent shoots. In this review, a synopsis is presented about how plants, especially macrophytes, respond to heavy metal stress; and we propose potential roles that phytohormones can play in the alleviation of metal toxicity in the aquatic environment. We focus on the uptake, translocation, and accumulation mechanisms of heavy metals in organs of macrophytes and give examples of how phytohormones interact with plant defense systems under heavy metal exposure. We advocate for a more in-depth understanding of these processes to inform more effective metal remediation techniques from metal-polluted water bodies. Environ Toxicol Chem 2021;40:7–22. © 2020 SETAC Abstract Heavy metal absorption, translocation, and accumulation within organs of aquatic plants and the changes in phytohormone signaling in response to heavy metal exposure. ABA = abscisic acid; BR = brassinosteroid; CK = cytokinin; ETH = ethylene; GA = gibberellin; JA = jasmonic acid; PA = polyamine; SA = salicylic acid.

Nod factors (NFs) are lipochitooligosaccharidic signal molecules produced by rhizobia, which play a key role in the rhizobium-legume symbiotic interaction. In this study, we analyzed the gene expression reprogramming induced by purified NF (4 and 24 h of treatment) in the root epidermis of the model legume Medicago truncatula. Tissue-specific transcriptome analysis was achieved by laser-capture microdissection coupled to high-depth RNA sequencing. The expression of 17,191 genes was detected in the epidermis, among which 1,070 were found to be regulated by NF addition, including previously characterized NF-induced marker genes. Many genes exhibited strong levels of transcriptional activation, sometimes only transiently at 4 h, indicating highly dynamic regulation. Expression reprogramming affected a variety of cellular processes, including perception, signaling, regulation of gene expression, as well as cell wall, cytoskeleton, transport, metabolism, and defense, with numerous NF-induced genes never identified before. Strikingly, early epidermal activation of cytokinin (CK) pathways was indicated, based on the induction of CK metabolic and signaling genes, including the CRE1 receptor essential to promote nodulation. These transcriptional activations were independently validated using promoter:β-glucuronidase fusions with the MtCRE1CK receptor gene and a CK response reporter (TWO COMPONENT SIGNALING SENSOR NEW). A CK pretreatment reduced the NF induction of the EARLY NODULIN11 (ENOD11) symbiotic marker, while a CK-degrading enzyme (CYTOKININ OXIDASE/DEHYDROGENASE3) ectopically expressed in the root epidermis led to increased NF induction of ENOD11 and nodulation. Therefore, CK may play both positive and negative roles in M. truncatula nodulation.

Cytokinins (CKs) are key phytohormones that not only regulate plant growth and development but also mediate plant tolerance to drought stress. Recent advances in genome-wide association studies coupled with in planta characterization have opened new avenues to investigate the drought-responsive expression of CK metabolic and signaling genes, as well as their functions in plant adaptation to drought. Under water deficit, CK signaling has evolved as an inter-cellular communication network which is essential to crosstalk with other types of phytohormones and their regulating pathways in mediating plant stress response. In this review, we revise the current understanding of CK involvement in drought stress tolerance. Particularly, a genetic framework for CK signaling and CK crosstalk with abscisic acid (ABA) in the precise monitoring of drought responses is proposed. In addition, the potential of endogenous CK alteration in crops towards developing drought-tolerant crops is also discussed.

Plants produce cytokinin (CK) hormones for controlling key developmental processes like source/sink distribution, cell division or programmed cell-death. Some plant pathogens have been shown to produce CKs but the function of this mimicry production by non-tumor inducing pathogens, has yet to be established. Here we identify a gene required for CK biosynthesis, CKS1, in the rice blast fungus Magnaporthe oryzae. The fungal-secreted CKs are likely perceived by the plant during infection since the transcriptional regulation of rice CK-responsive genes is altered in plants infected by the mutants in which CKS1 gene was deleted. Although cks1 mutants showed normal in vitro growth and development, they were severely affected for in planta growth and virulence. Moreover, we showed that the cks1 mutant triggered enhanced induction of plant defenses as manifested by an elevated oxidative burst and expression of defense-related markers. In addition, the contents of sugars and key amino acids for fungal growth were altered in and around the infection site by the cks1 mutant in a different manner than by the control strain. These results suggest that fungal-derived CKs are key effectors required for dampening host defenses and affecting sugar and amino acid distribution in and around the infection site. The role of plant-like hormonal compounds produced by fungal pathogens during infection has not been elucidated. Here we identified a conserved gene in most fungi, required for cytokinin production by the rice blast fungus and for its full virulence. Fungal-derived cytokinins are likely potent inhibitors of plant immunity. They are also needed to maintain elevated sugar contents at the site of infection and to drain or consume essential amino acids at, and around, the infection site. Thus, cytokinins represent the second example, after the bacterially-produced coronatine, of plant hormones hijacked by pathogens to successfully invade plant tissues. These findings also suggest that this invasion strategy could be widely conserved among fungi.

Background: Symbiotic Methylobacterium strains comprise a significant part of plant microbiomes. Their presence enhances plant productivity and stress resistance, prompting classification of these strains as plant growth-promoting bacteria (PGPB). Methylobacteria can synthesize unusually high levels of plant hormones, called cytokinins (CKs), including the most active form, trans-Zeatin (tZ). Results: This study provides a comprehensive inventory of 46 representatives of Methylobacterium genus with respect to phytohormone production in vitro, including 16 CK forms, abscisic acid (ABA) and indole-3-acetic acid (IAA). High performance-liquid chromatography—tandem mass spectrometry (HPLC–MS/MS) analyses revealed varying abilities of Methylobacterium strains to secrete phytohormones that ranged from 5.09 to 191.47 pmol mL⁻¹ for total CKs, and 0.46 to 82.16 pmol mL⁻¹ for tZ. Results indicate that reduced methanol availability, the sole carbon source for bacteria in the medium, stimulates CK secretion by Methylobacterium. Additionally, select strains were able to transform L-tryptophan into IAA while no ABA production was detected. Conclusions: To better understand features of CKs in plants, this study uncovers CK profiles of Methylobacterium that are instrumental in microbe selection for effective biofertilizer formulations.