Phytohormone biosynthesis and signaling pathways of mosses.

KEY MESSAGE: Most known phytohormones regulate moss development. We present a comprehensive view of the synthesis and signaling pathways for the most investigated of these compounds in mosses, focusing on the model Physcomitrium patens. The last 50 years of research have shown that most of the known phytohormones are synthesized by the model moss Physcomitrium patens (formerly Physcomitrella patens) and regulate its development, in interaction with responses to biotic and abiotic stresses. Biosynthesis and signaling pathways are best described in P. patens for the three classical hormones auxins, cytokinins and abscisic acid. Furthermore, their roles in almost all steps of development, from early filament growth to gametophore development and sexual reproduction, have been the focus of much research effort over the years. Evidence of hormonal roles exist for ethylene and for CLE signaling peptides, as well as for salicylic acid, although their possible effects on development remain unclear. Production of brassinosteroids by P. patens is still debated, and modes of action for these compounds are even less known. Gibberellin biosynthesis and signaling may have been lost in P. patens, while gibberellin precursors such as ent-kaurene derivatives could be used as signals in a yet to discover pathway. As for jasmonic acid, it is not used per se as a hormone in P. patens, but its precursor OPDA appears to play a corresponding role in defense against abiotic stress. We have tried to gather a comprehensive view of the biosynthesis and signaling pathways for all these compounds in mosses, without forgetting strigolactones, the last class of plant hormones to be reported. Study of the strigolactone response in P. patens points to a novel signaling compound, the KAI2-ligand, which was likely employed as a hormone prior to land plant emergence.

Physico-Chemical Properties of Inorganic NPs Influence the Absorption Rate of Aquatic Mosses Reducing Cytotoxicity on Intestinal Epithelial Barrier Model.

Noble metals nanoparticles (NPs) and metal oxide NPs are widely used in different fields of application and commercial products, exposing living organisms to their potential adverse effects. Recent evidences suggest their presence in the aquifers water and consequently in drinking water. In this work, we have carefully synthesized four types of NPs, namely, silver and gold NPs (Ag NPs and Au NPs) and silica and titanium dioxide NPs (SiO2 NPs and TiO2 NPs) having a similar size and negatively charged surfaces. The synthesis of Ag NPs and Au NPs was carried out by colloidal route using silver nitrate (AgNO3) and tetrachloroauric (III) acid (HAuCl4) while SiO2 NPs and TiO2 NPs were achieved by ternary microemulsion and sol-gel routes, respectively. Once the characterization of NPs was carried out in order to assess their physico-chemical properties, their impact on living cells was studied. We used the human colorectal adenocarcinoma cells (Caco-2), known as the best representative intestinal epithelial barrier model to understand the effects triggered by NPs through ingestion. Then, we moved to explore how water contamination caused by NPs can be lowered by the ability of three species of aquatic moss, namely, Leptodictyum riparium, Vesicularia ferriei, and Taxiphyllum barbieri, to absorb them. The experiments were conducted using two concentrations of NPs (100 µM and 500 Μm as metal content) and two time points (24 h and 48 h), showing a capture rate dependent on the moss species and NPs type. Then, the selected moss species, able to actively capture NPs, appear as a powerful tool capable to purify water from nanostructured materials, and then, to reduce the toxicity associated to the ingestion of contaminated drinking water.

Assessment of cultured media for desert moss crust by physiological responses.

The physiological responses of desert moss crusts under four artificial media (Beneck, Part, BG11, and Hogland) were investigated to evaluate the function of culture media during different culture periods. The results showed that the value of malondialdehyde (MDA) was at a maximum at 11d, on the contrary, chlorophyll-a, soluble protein, and soluble sugar were at a minimum. As the time increased, the value of MDA and soluble protein decreased faster in the Hogland, while the value of chlorophyll-a and soluble sugar increased. At the end of the culture period, the value of chlorophyll-a and soluble sugar was at a maximum in the Hogland, while the value of MDA and soluble protein was at a minimum. The results suggested that the Hogland medium had a promoting effect on the growth of desert moss crusts. The selected artificial cultivation medium towards wider and larger scale field applications of cultural desert biocrust was widely anticipated.

Effects of wood distillate (pyroligneous acid) on sensitive bioindicators (lichen and moss).

Wood distillate (pyroligneous acid) can be successfully applied in agriculture to increase crop quality and productivity with a lower risk for the environment respect to synthetic chemical herbicides, pesticides or fertilizers. However, the effects of wood distillate on the environment and biota are still under investigation, depending on biological attributes of potentially influenced organisms. The potential toxicological effects of wood distillate on sensitive non-target organisms, lichens and mosses, are studied for the first time. The physiological parameters (chlorophyll a fluorescence emission FV/FM and PI(ABS), chlorophyll content, spectral reflectance, antioxidant power, and dehydrogenase activity) and eventual bioaccumulation of selected elements (As, Ba, Cd, Cr, Cu, Fe, Ni, Pb, Zn) were investigated in the lichen Xanthoria parietina and the moss Hypnum cupressiforme after short-term treatments over a range of wood distillate solutions (1:300, 1:500, 1:700) to detect potential early stress responses. Overall, the lichen did not show changes after the treatments, while in the moss wood distillate caused only modest alterations in FV/FM and PI(ABS) and progressive increasing of antioxidant activity according to the dose supplied. The bioaccumulation of toxic elements was low and did not show any pattern of uptake with increasing concentrations of wood distillate.

Effects of environmental factors and heavy metals on the vertical distribution of bryophytes in a sinkhole environment.

A sinkhole ecosystem, as a refuge for plant diversity, has been subjected to intensive exploitation, leading to ecosystem destruction of sinkholes in China. Understanding the responses of bryophyte distribution to destruction of the sinkhole environment are crucial to implementing protection measures for bryophyte diversity. Haolong sinkhole in Guangxi Zhuang Autonomous Region of China, the third largest sinkhole in the world, was selected as the study area. The Wilson Shmida index was used to analyse bryophyte species diversity; a Generalized Linear Model (GLM) was used to reveal species vertical distribution of bryophytes, the Single and Multiple Species Distribution Models (SSDM, MSDM) were used for analysis of the relationship between bryophyte species distribution, environmental factors and heavy metals. A total of 183 species from 74 genera in 36 families of bryophytes were collected from Haolong sinkhole, of which 26 species are endemic to China. Bryophyte species diversity was ranked in the order: agricultural section < forest section < grassland. In the vertical direction, bryophyte distribution was divided into point, disjunctive and continuous distributions using the GLM. The SSMA and MSDM indicated that bryophyte species of each of these three distributions can be divided into a temperature-slope zone, light-depth-pH-humidity zone, Pb (B)-Hg (B) zone and mixed heavy metals zone according to the effect of environmental factors and heavy metals such as As. Environmental factors or heavy metals, such as As, in Haolong sinkhole effectively cooperate in bryophyte distribution. An effective way to protect bryophyte diversity, in particular species endemic to China in the sinkhole environment, is through education and involvement of the local villagers to minimize further damage to the sinkhole environment.

Perspective of mitigating atmospheric heavy metal pollution: using mosses as biomonitoring and indicator organism.

Mosses were proved as an ideal and reliable biomonitor as well as an indicator of atmospheric trace metal pollution. They are used as model indicator species of air pollution since long back due to their simple structure, genetic diversity, totipotency, rapid colony-forming ability, and high metal resistance behavior. Bryomonitoring technique is gradually being popularized as an economically viable procedure for estimating the degrees of environmental health and evaluating the toxic pollutants in biosphere. Thus, in the present scenario, many parts of the world use these organisms for monitoring the air pollution. This article describes an overview of the relationship of terrestrial mosses with trace metals with respect to their uptake, accumulation, and toxification as well as detoxification and tolerance mechanisms. The review article explicitly expresses the caliber of the cryptogamic mosses in establishing the pristine environment around the world. It also highlights the underpinning mechanisms and potential for future research directions. We have referred more than 250 articles, which deals with the assessment and impact of different heavy metals on 52 numbers of different moss species belongs to different climatic zones. The present review covers the research work in this area carried out worldwide since 1965.

Long-term exposure to atmospheric metals assessed by mosses and mortality in France.

BACKGROUND: Long-term exposure to air pollution affects health, but little is known about exposure to atmospheric metals. Estimating exposure to atmospheric metals across large spatial areas remains challenging. Metal concentrations in mosses could constitute a useful proxy. Here, we linked moss biomonitoring and epidemiological data to investigate the associations between long-term exposure to metals and mortality. METHODS: We modelled and mapped 13 atmospheric metals from a 20-year national moss biomonitoring program to derive exposure estimates across France. In the population-based Gazel cohort, we included 11,382 participants from low to intermediate population density areas and assigned modelled metals to their residential addresses. We distinguished between airborne metals that are primarily of natural origin and those primarily of anthropogenic origin. Associations were estimated between exposure to metals and mortality (natural-cause, cardiovascular and respiratory), using Cox models, with confounder adjustment at individual level. FINDINGS: Between 1996 and 2017, there were 1313 deaths in the cohort (including 181 cardiovascular and 33 respiratory). Exposure to the anthropogenic metals was associated with an increased risk of natural-cause mortality (hazard ratio of 1.16 [1.08-1.24] per interquartile range of exposure), while metals from natural sources were not. INTERPRETATION: Some atmospheric anthropogenic metals may be associated with excess mortality - even in areas with relatively low levels of exposure to air pollution. Consistent with the previous literature, our findings support the use of moss biomonitoring as a tool to assess health effects of air pollution exposure at individual level.

Abscisic acid, cold and salt stimulate conserved metabolic regulation in the moss Physcomitrella patens.

Salt and cold are major abiotic stresses that have adverse effects on plant growth and development. To cope with these stresses and their detrimental effects plants have evolved several metabolic, biochemical and physiological processes that are mainly triggered and mediated by the plant hormone abscisic acid (ABA). To elucidate the metabolic responses of the moss Physcomitrella patens, which serves as a model plant for abiotic stress adaptation, we performed GC-MS-based metabolic profiling of plants challenged for 5 and 28 h with either salt, cold or ABA. Our results indicate significant changes in the accumulation of several sugars including maltose, isomaltose and trehalose, amino acids including arginine, histidine, ornithine, tryptophan and tyrosine, and organic acids mainly citric acid and malonic acid. The metabolic responses provoked by ABA, cold and salt show considerable similarities. The accumulation of certain metabolites positively correlates with gene expression data whereas some metabolites do not show correlation with cognate transcript abundance. To place our results into an evolutionary context we compared the ABA- and stress-induced metabolic changes in moss to available metabolic profiles of the seed plant Arabidopsis thaliana. We detected considerable conservation between the species, indicating early evolution of stress-associated metabolic adaptations that probably occurred at the plant water-to-land transition.

Mechanisms of drought-induced dissipation of excitation energy in sun- and shade-adapted drought-tolerant mosses studied by fluorescence yield change and global and target analysis of fluorescence decay kinetics.

Some mosses stay green and survive long even under desiccation. Dissipation mechanisms of excess excitation energy were studied in two drought-tolerant moss species adapted to contrasting niches: shade-adapted Rhytidiadelphus squarrosus and sun-adapted Rhytidium rugosum in the same family. (1) Under wet conditions, a light-induced nonphotochemical quenching (NPQ) mechanism decreased the yield of photosystem II (PSII) fluorescence in both species. The NPQ extent saturated at a lower illumination intensity in R. squarrosus, suggesting a larger PSII antenna size. (2) Desiccation reduced the fluorescence intensities giving significantly lower F 0 levels and shortened the overall fluorescence lifetimes in both R. squarrosus and R. rugosum, at room temperature. (3) At 77 K, desiccation strongly reduced the PSII fluorescence intensity. This reduction was smaller in R. squarrosus than in R. rugosum. (4) Global and target analysis indicated two different mechanisms of energy dissipation in PSII under desiccation: the energy dissipation to a desiccation-formed strong fluorescence quencher in the PSII core in sun-adapted R. rugosum (type-A quenching) and (5) the moderate energy dissipation in the light-harvesting complex/PSII in shade-adapted R. squarrosus (type-B quenching). The two mechanisms are consistent with the different ecological niches of the two mosses.

The effects of experimentally supplied lead nitrate on three common Mediterranean moss species.

We assess here, through an experimental simulation using lead nitrate, the response to lead deposition of three common Mediterranean bryophyte species in the family Pottiaceae. Five concentrations of lead nitrate (from 0 to 10-3 M) were sprayed for 4 months on plants belonging to Tortula muralis (reported as toxitolerant), Syntrichia ruralis (medium-tolerant), and Tortula subulata (less tolerant). The three species showed a remarkably high tolerance to lead nitrate, with a low incidence of damage even at concentrations as high as 10-4 M. The maximum concentration (10-3 M), although resulting eventually in serious damages in the gametophyte of the three species (high mortality rates in S. ruralis and T. subulata, or a significant percentage of damaged tissue in T. muralis), did not prevent the production of sporophytes in the two species with fertile samples (T. muralis and T. subulata). Growth parameters show limited value as bioindicators of lead deposition, as they only show clear effects at very high concentrations. Besides, we identified the existence of a lead exclusion strategy mediated by mucilage using histochemical analyses and scanning electron microscopy combined with energy-dispersive X-ray spectroscopy. This mechanism can hamper the usefulness of these mosses in quantitative estimation of lead deposition.

Diversity of strategies for escaping reactive oxygen species production within photosystem I among land plants: P700 oxidation system is prerequisite for alleviating photoinhibition in photosystem I.

In land plants, photosystem I (PSI) photoinhibition limits carbon fixation and causes growth defects. In addition, recovery from PSI photoinhibition takes much longer than PSII photoinhibition when the PSI core-complex is degraded by oxidative damage. Accordingly, PSI photoinhibition should be avoided in land plants, and land plants should have evolved mechanisms to prevent PSI photoinhibition. However, such protection mechanisms have not yet been identified, and it remains unclear whether all land plants suffer from PSI photoinhibition in the same way. In the present study, we focused on the susceptibility of PSI to photoinhibition and investigated whether mechanisms of preventing PSI photoinhibition varied among land plant species. To assess the susceptibility of PSI to photoinhibition, we used repetitive short-pulse (rSP) illumination, which specifically induces PSI photoinhibition. Subsequently, we found that land plants possess a wide variety of tolerance mechanisms against PSI photoinhibition. In particular, gymnosperms, ferns and mosses/liverworts exhibited higher tolerance to rSP illumination-induced PSI photoinhibition than angiosperms, and detailed analyses indicated that the tolerance of these groups could be partly attributed to flavodiiron proteins, which protected PSI from photoinhibition by oxidizing the PSI reaction center chlorophyll (P700) as an electron acceptor. Furthermore, we demonstrate, for the first time, that gymnosperms, ferns and mosses/liverworts possess a protection mechanism against photoinhibition of PSI that differs from that of angiosperms.

Arsenic in stream waters is bioaccumulated but neither biomagnified through food webs nor biodispersed to land.

Human activities such as mining have contributed substantially to the increase of metals in aquatic environments worldwide. These metals are bioaccumulated by aquatic organisms and can be biomagnified along trophic webs. The dispersal of contaminants from water to land has been little investigated, even though most aquatic invertebrates in streams have aerial stages. We used field and laboratory approaches to investigate the effects of arsenic pollution on stream invertebrate assemblages, and its bioaccumulation, biomagnification and trophic transfer from aquatic to terrestrial environments by emergent insects. We conducted the study in an arsenic-impacted stream (40µgL-1 As at the most polluted site) and a reference stream (0.3µgL-1 As). Invertebrate abundance and richness were lowest at the most impacted site. Arsenic in biofilm and in invertebrates increased with the arsenic content in the water. The highest arsenic accumulators were bryophytes (1760µgg-1), followed by the biofilm (449µgg-1) and shredder invertebrates (313µgg-1); predators had the lowest arsenic concentration. Insects emerging from water and spiders along streambanks sampled from the reference and the impacted stream did not differ in their body arsenic concentrations. In the laboratory, the shredder Sericostoma vittatum had reduced feeding rates when exposed to water from the impacted stream in comparison with the reference stream (15.6 vs. 19.0mg leaves mg body mass-1 day-1; p<0.05), but they grew faster in the polluted water (0.16±0.04% day-1vs. 0.05±0.01% day-1, p<0.05). S. vittatum exposed to contaminated stream water accumulated arsenic from food, not through contact with water. We concluded that although arsenic is bioaccumulated, mainly by food ingestion, it is not biomagnified through food webs and is not transported from the aquatic to terrestrial environment when insects leave the stream water.

Mercury in the ecosystem of Admiralty Bay, King George Island, Antarctica: Occurrence and trophic distribution.

Mercury (Hg) can reach the environment through natural and human-related sources, threatening ecosystems all over the planet due to its well known deleterious effects. Therefore, Antarctic trophic webs, despite being relatively isolated, are not exempt of its influence. To evaluate Hg concentrations in an Antarctic ecosystem, different tissues from 2 species of invertebrates, 2 of fish, 8 of birds, 4 of pinnipeds and at least 5 of vegetation were investigated (n=176). For animals, values ranged from 0.018 to 48.7µgg-1 dw (whole Antarctic krill and Antarctic Fur Seal liver). They were generally correlated to trophic position (assessed by δ15N and δ13C) but also to cephalopods and myctophids consumption. For vegetation, values ranged from 0.014 to 0.227µgg-1 dw (Colobanthus quitensis and an unidentified lichen), with lichens presenting significantly higher values than mosses, likely due to year-round exposure and absorption of animal derived organic matter, as hypothesized by literature.

Multivariate extraction of dominant geochemical markers for deposition of 69 elements in the Bregalnica River basin, Republic of Macedonia (moss biomonitoring).

Atmospheric deposition was investigated using the terrestrial moss species Hypnum cupressiforme (Hedw.) and Homolothecium lutescens (Hedw.) in the Bregalnica River basin, Republic of Macedonia. Long-term emission occurs in this area due to the hydrothermal exploitation of Pb-Zn deposits (Sasa and Zletovo mines) and copper ore exploitation and floatation (Bucim mine). Determination of the chemical elements was conducted using atomic emission spectrometry with inductively coupled plasma (ICP-AES) and mass spectrometry with inductively coupled plasma (ICP-MS). A combination of multivariate techniques (PCA, FA and CA) was applied for data processing and identification of element association with lithogenic/anthropogenic origin. Seven dominant factors were extracted from the total of 69 analysed elements. Spatial distribution maps were constructed for the determination and localisation of smaller areas with higher contents of certain anthropogenic elements. In this way, the influences of selected human activities on local air pollution can be determined. The summarised data show quantification of the element distributions. This not only allows the determination of the distribution of hazardous elements but also presents complete characterisation of element deposition in the environs of mines.

Trace element concentrations in the moss Hypnum cupressiforme growing in a presumably unpolluted area.

In this study we determined the concentrations of As, Cd, Hg, Ni and Pb in samples of the moss Hypnum cupressiforme collected during 5 different sampling surveys (2006-2014) in a presumably unpolluted area in northern Spain (25 sampling sites). We then applied factor analysis (FA) to the data to explore the factors underlying the spatial and temporal variability in the concentrations. The percentage of variance explained by the FA ranged between 34 and 98%, and was usually higher than 70%. The FA yielded 5 factors that explained the variance in the concentrations of Cd, As, Hg and Pb in all sampling surveys and also a single factor that explained the variance in Hg and Pb concentrations in 2006. Although the lack of obvious sources of pollution in the study region (at least for the elements considered) suggests that most elements (except perhaps Ni) probably originated from long-range atmospheric transport, this would not explain the results of the FA. We suggest that rather than being due to the origin of the pollutants (as frequently assumed), the spatio-temporal variability in the concentrations of these elements is probably determined by a series of other factors: the physicochemical characteristics of the pollutants and of the moss binding surfaces, physiological processes (e.g. moss growth), and the characteristics of the sampling sites (e.g. vegetation cover, elevation, slope, aspect). We therefore conclude that the assumption that variations in element concentrations in moss tissues are due to the origin of the pollutants is an oversimplification that leads to erroneous interpretation of the results of biomonitoring studies with terrestrial mosses.

Changes in bryophyte and lichen communities on Scots pines along an alkaline dust pollution gradient.

Dust pollution can cause a significant damage of environment and endanger human health. Our study aimed to investigate epiphytic lichens and bryophytes in relation to long-term alkaline dust pollution and provide new insights into the bioindicators of dust pollution. We measured the bark pH of Scots pines and the species richness and cover of two cryptogam groups in 32 sample plots in the vicinity of limestone quarries (up to ca. 3 km) in northern Estonia. The bark pH decreased gradually with increasing distance from quarries. We recorded the changes in natural epiphytic communities, resulting in diversified artificial communities on pines near the pollution source; the distance over 2 km from the quarries was sufficient to re-establish the normal acidity of the bark and natural communities of both lichens and bryophytes. The cover of lichens and the number of bryophytes are a more promising indicator of environmental conditions than individual species occurrence. We confirmed previously proposed and suggested new bioindicator species of dust pollution (e.g., Lecidella elaeochroma, Opegrapha varia, Schistidium apocarpum). Limestone quarrying activity revealed a "parapositive" impact on cryptogamic communities, meaning that quarrying might, besides disturbances of natural communities, temporarily contribute to the distribution of locally rare species.

The copper spoil heap Knappenberg, Austria, as a model for metal habitats - Vegetation, substrate and contamination.

Historic mining in the Eastern Alps has left us with a legacy of numerous spoil heaps hosting specific, metal tolerant vegetation. Such habitats are characterized by elevated concentrations of toxic elements but also by high irradiation, a poorly developed substrate or extreme pH of the soil. This study investigates the distribution of vascular plants, mosses and lichens on a copper spoil heap on the ore bearing Knappenberg formed by Prebichl Layers and Werfener Schist in Lower Austria. It serves as a model for discriminating between various ecological traits and their effects on vegetation. Five distinct clusters were distinguished: (1) The bare, metal rich Central Spoil Heap was only colonised by highly resistant specialists. (2) The Northern and (3) Southern Peripheries contained less copper; the contrasting vegetation was best explained by the different microclimate. (4) A forest over acidic bedrock hosted a vegetation overlapping with the periphery of the spoil heap. (5) A forest over calcareous bedrock was similar to the spoil heap with regard to pH and humus content but hosted a vegetation differing strongly to all other habitats. Among the multiple toxic elements at the spoil heap, only Cu seems to exert a crucial influence on the vegetation pattern. Besides metal concentrations, irradiation, humidity, humus, pH and grain size distribution are important for the establishment of a metal tolerant vegetation. The difference between the species poor Northern and the diverse Southern Periphery can be explained by the microclimate rather than by the substrate. All plant species penetrating from the forest into the periphery of the spoil heap originate from the acidic but not from the calcareous bedrock.

Using devitalized moss for active biomonitoring of water pollution.

This paper presents the results of an experiment carried out for the first time in situ to select a treatment to devitalize mosses for use in active biomonitoring of water pollution. Three devitalizing treatments for the aquatic moss Fontinalis antipyretica were tested (i.e. oven-drying at 100 °C, oven-drying with a 50-80-100 °C temperature ramp, and boiling in water), and the effects of these on loss of material during exposure of the transplants and on the accumulation of different heavy metals and metalloids were determined. The suitability of using devitalized samples of the terrestrial moss Sphagnum denticulatum to biomonitor aquatic environments was also tested. The structure of mosses was altered in different ways by the devitalizing treatments. Devitalization by boiling water led to significantly less loss of material (p < 0.01) than the oven-drying treatments. However, devitalization by oven-drying with a temperature ramp yielded more stable results in relation to both loss of material and accumulation of elements. With the aim of standardizing the moss bag technique, the use of F. antipyretica devitalized by oven-drying with a temperature ramp is recommended, rather than other devitalization treatments or use of S. denticulatum.

The nitric oxide production in the moss Physcomitrella patens is mediated by nitrate reductase.

During the last 20 years multiple roles of the nitric oxide gas (•NO) have been uncovered in plant growth, development and many physiological processes. In seed plants the enzymatic synthesis of •NO is mediated by a nitric oxide synthase (NOS)-like activity performed by a still unknown enzyme(s) and nitrate reductase (NR). In green algae the •NO production has been linked only to NR activity, although a NOS gene was reported for Ostreococcus tauri and O. lucimarinus, no other Viridiplantae species has such gene. As there is no information about •NO synthesis neither for non-vascular plants nor for non-seed vascular plants, the interesting question regarding the evolution of the enzymatic •NO production systems during land plant natural history remains open. To address this issue the endogenous •NO production by protonema was demonstrated using Electron Paramagnetic Resonance (EPR). The •NO signal was almost eliminated in plants treated with sodium tungstate, which also reduced the NR activity, demonstrating that in P. patens NR activity is the main source for •NO production. The analysis with confocal laser scanning microscopy (CLSM) confirmed endogenous NO production and showed that •NO signal is accumulated in the cytoplasm of protonema cells. The results presented here show for the first time the •NO production in a non-vascular plant and demonstrate that the NR-dependent enzymatic synthesis of •NO is common for embryophytes and green algae.