Bamboos stand out among other tall plants in being able to generate positive pressure in the xylem at night, pushing water up to the leaves and causing drops to fall from leaf tips as guttation that can amount to a steady nocturnal \'bamboo rain\'. The location and mechanism of nocturnal pressure generation in bamboos are unknown, as are the benefits for the plants. We conducted a study on the tall tropical bamboo species Bambusa oldhamii (giant timber bamboo) growing outdoors in southern California under full irrigation to determine where in the plant the nocturnal pressure is generated, when it rises in the evening, and when it dissipates in the morning. We hypothesized that the build-up of positive pressure would be triggered by the cessation of transpiration-driven sap flow and that resumption of sap flow in the morning would cause the pressure to dissipate. Nocturnal pressure was observed in mature stems and rhizomes, but never in roots. The pressure was episodic and associated with stem swelling and was usually, but not always, higher in rhizomes and basal stems than in stems at greater height. Time series analyses revealed that dry atmospheric conditions were followed by lower nocturnal pressure and rainfall events by higher stem pressure. Nocturnal pressure was unrelated to sap flow and even was generated for a short time in isolated stem pieces placed in water. We conclude that nocturnal pressure in bamboo is not \'root pressure\' but is generated in the pseudo-woody rhizomes and stems. It is unrelated to the presence or absence of sap flow and therefore must be created outside of vessels, such as in phloem, parenchyma, or fibres. It is unlikely to be a drought adaptation and may benefit the plants by maximizing stem water storage for daytime transpiration or by transporting nutrients to the leaves.

Solar radiation triggers atmospheric nitrous acid (HONO) photolysis, producing OH radicals, thereby accelerating photochemical reactions, leading to severe secondary pollution formation. Missing daytime sources were detected in the extensive HONO budget studies carried out in the past. In the rural North China Plain, some studies attributed those to soil emissions and more recent studies to dew evaporation. To investigate the contributions of these two processes to HONO temporal variations and unknown production rates in rural areas, HONO and related field observations obtained at the Gucheng Agricultural and Ecological Meteorological Station during spring and autumn were thoroughly analyzed. Morning peaks in HONO frequently occurred simultaneously with those of ammonia (NH3) and water vapor both during spring and autumn, which were mostly caused by dew and guttation water evaporation. In spring, the unknown HONO production rate revealed pronounced afternoon peaks exceeding those in the morning. In autumn, however, the afternoon peak was barely detectable compared to the morning peak. The unknown afternoon HONO production rates were attributed to soil emissions due to their good relationship to soil temperatures, while NH3 soil emissions were not as distinctive as dew emissions. Overall, the relative daytime contribution of dew emissions was higher during autumn, while soil emissions dominated during spring. Nevertheless, dew emission remained the most dominant contributor to morning time HONO emissions in both seasons, thus being responsible for the initiation of daytime OH radical formation and activation of photochemical reactions, while soil emissions further maintained HONO and associated OH radial formation rates at a high level, especially during spring. Future studies need to thoroughly investigate the influencing factors of dew and soil emissions and establish their relationship to HONO emission rates, form reasonable parameterizations for regional and global models, and improve current underestimations in modeled atmospheric oxidation capacity.

The unknown daytime source of HONO has been extensively investigated due to unexplained atmospheric oxidation capacity and current modelling bias, especially during cold seasons. In this study, abrupt morning increases in atmospheric HONO at a rural site in the North China Plain (NCP) were observed almost on daily basis, which were closely linked to simultaneous rises in atmospheric water vapor content and NH3 concentrations. Dew and guttation water formation was frequently observed on wheat leaves, from which water samples were taken and chemically analyzed for the first time. Results confirmed that such natural processes likely governed the daily nighttime deposition and daytime release of HONO and NH3, which have not been considered in the numerous HONO budget studies investigating its large missing daytime source in the NCP. The dissolved HONO and NH3 in leaf surface water droplets reached 1.4 and 23 mg L-1 during the morning on average, resulting in averaged atmospheric HONO and NH3 increases of 0.89 ± 0.61 and 43.7 ± 29.3 ppb during morning hours, with relative increases of 186 ± 212 % and 233 ± 252 %, respectively. The high atmospheric oxidation capacity contained within HONO was stored in near surface liquid water (such as dew, guttation and soil surface water) during nighttime, which prevented its atmospheric dispersion after sunset and protected it from photodissociation during early morning hours. HONO was released in a blast during later hours with stronger solar radiation, which triggered and then accelerated daytime photochemistry through the rapid photolysis of HONO and subsequent OH production, especially under high RH conditions, forming severe secondary gaseous and particulate pollution. Results of this study demonstrate that global ecosystems might play significant roles in atmospheric photochemistry through nighttime dew formation and guttation processes.

Plants prevent disease by passively and actively protecting potential entry routes against invading microbes. For example, the plant immune system actively guards roots, wounds, and stomata. How plants prevent vascular disease upon bacterial entry via guttation fluids excreted from specialized glands at the leaf margin remains largely unknown. These so-called hydathodes release xylem sap when root pressure is too high. By studying hydathode colonization by both hydathode-adapted (Xanthomonas campestris pv. campestris) and non-adapted pathogenic bacteria (Pseudomonas syringae pv. tomato) in immunocompromised Arabidopsis mutants, we show that the immune hubs BAK1 and EDS1-PAD4-ADR1 restrict bacterial multiplication in hydathodes. Both immune hubs effectively confine bacterial pathogens to hydathodes and lower the number of successful escape events of an hydathode-adapted pathogen toward the xylem. A second layer of defense, which is dependent on the plant hormones\' pipecolic acid and to a lesser extent on salicylic acid, reduces the vascular spread of the pathogen. Thus, besides glands, hydathodes represent a potent first line of defense against leaf-invading microbes.

Guttation, the formation of exudation water, is widespread among plants and fungi, yet the underlying mechanisms remain largely unknown. We describe the conditions for inducing guttation in sporangiophores of the mucoracean fungus, Phycomyces blakesleeanus. Cultivation on peptone-enriched potato dextrose agar elicits vigorous guttation mainly below the apical growing zone, while sporangiophores raised on a glucose-mineral medium manifest only moderate guttation. Mycelia do not guttate irrespective of the employed media. The topology of guttation droplets allows identifying the non-growing part of the sporangiophore as a guttation zone, which responds to humidity and medium composition in ways that become relevant for turgor homeostasis and thus the sensor physiology of the growing zone. Apparently, the entire sporangiophore, rather than exclusively the growing zone, participates in signal reception and integration to generate a common growth output. Exogenous auxin applied to the growing zones elicits two correlated responses: (i) formation of guttation droplets in the growing and transition zones below the sporangium and (ii) a diminution of the growth rate. In sporangiophore populations, guttation-induction by exogenous control buffer occurs at low frequencies; the bias for guttation increases with increasing auxin concentration. Synthetic auxins and the transport inhibitor NPA suppress guttation completely, but leave growth rates largely unaffected. Mutants C2 carA and C148 carA madC display higher sensitivities for auxin-induced guttation compared to wild type. A working model for guttation includes aquaporins and mechanosensitive ion channels that we identified in Phycomyces by sequence domain searches.

Ferns are the second largest group of vascular plants and are distributed nearly worldwide. Although ferns have been integrated into some comparative ecological studies focusing on hydathodes, there is a considerable gap in our understanding of the functional anatomy of these secretory tissues that are found on the vein endings of many fern leaves. In this study, we aimed to investigate the phylogenetic distribution, structure and function of fern hydathodes.
We performed a global review on fern hydathodes and their phylogenetic distribution, carried out an ancestral character state reconstruction, and studied the structure, guttation and elemental composition of salt residues of eight species, and the diurnal patterns of xylem pressure of two species.
Hydathodes are known from 1189 fern species, 92 genera and 19 families of 2 orders, Equisetales and Polypodiales. Stochastic character mapping indicated multiple gains and losses of hydathodes at the genus level, occurring especially during the last 50 million years of fern evolution. Hydathodes were located on the adaxial leaf surface and characterized by a cytoplasm-rich, pore-free epidermis, and became functional for several weeks after nearly complete leaf expansion. In two species, positive xylem pressure built up at night, potentially facilitating guttation. Guttation fluid was rich in Ca and often Si, but also contained P, Mg, Na and Al.
Stochastic character mapping and the structural and functional diversity of hydathodes indicate multiple origins, and their presence/absence in closely related taxa implies secondary losses during fern evolution. Positive xylem pressure and high air humidity play an important role as drivers of guttation. Hydathodes may contribute to the regulation of leaf nutrient stoichiometry by the release of excessive compounds and minerals other than waste products, but the presence of essential chemical elements in salt residues also indicates possible leakage.

Guttation is a common feature of cucumber leaves under high relative humidity conditions; however, little is known about the role of guttation in the transmission of Pseudomonas amygdali pv. lachrymans, which is the pathogen of cucumber angular leaf spot disease. In this study, experimental evidence for the transmission of P. amygdali pv. lachrymans inside cucumber plants and through guttation was provided, and the results proved that P. amygdali pv. lachrymans can be transmitted from the bottom leaf to the upper leaves inside the plant and excreted from the upper leaves through guttation. After that, the third leaf of cucumber was inoculated with P. amygdali pv. lachrymans bacterial suspension, P. amygdali pv. lachrymans was detected on the fifth leaf, the petiole, and the stem and in guttation drops. Healthy cucumber seedlings were infected by P. amygdali pv. lachrymans in the guttation droplets, indicating that guttation fluids containing P. amygdali pv. lachrymans could become a potential source of secondary infection. The results from this study verified the hypothesis that guttation is a potential route for P. amygdali pv. lachrymans excretion from cucumber plants and may be a source of secondary transmission under high relative humidity.

Production of guttation droplets in the cultures of Colletotrichum falcatum was often noticed. Hence, formation of guttation droplets, exudation of toxin in guttation, and guttation droplet chemical composition were assessed to find the ecological importance of guttation. Among the tested culture media, more guttation was noticed in oat meal agar medium and host extracts broth medium. Assessing the effect of five different carbon sources on guttation of C. falcatum showed that among solid media, sparse guttation droplets were noticed in glucose-, fructose-, and starch-amended media, whereas in broth cultures except fructose-amended media no other carbon sources induced guttation in mycelium. The guttation phenomenon was noticed only in sporulating mycelia, and it also followed a specific pattern in connection with sporulation. Bioassay of guttation exudates on sugarcane leaves produced pronounced necrotic lesions with large, yellow halo similar to positive control, thus confirming the presence of toxin. The guttation fluid initiated browning symptom on sugarcane leaves at a concentration of 1000 ppm and produced pronounced necrotic lesions of 3.7 cm at 10 000 ppm. The toxicity of crude fraction was thermostable up to 60 C. The gas chromatography-mass spectrometry (GC-MS) analysis of crude guttation droplets identified the presence of 24 compounds belonging to fatty acids, derivatives of fatty acids, flavonoids, flavonoid glycosides, sterols, quinone derivatives, amides, and glycol ethers, and among them fatty acids were dominant. The two compounds, viz., hexadecanoic acid methyl ester and 9-octadecenoic acid (Z) methyl ester, were abundant (18.14% and 17.98%, respectively) and identified as probable toxic compounds.

The secretion of metabolites in guttation droplets by indoor moulds is not well documented. This study demonstrates the guttation of metabolites by actively growing common indoor moulds. Old and fresh biomasses of indoor isolates of Aspergillus versicolor, Chaetomium globosum, Penicillium expansum, Trichoderma atroviride, T. trixiae, Rhizopus sp. and Stachybotrys sp. were compared. Metabolic activity indicated by viability staining and guttation of liquid droplets detected in young (<3 weeks old) biomass were absent in old (>6 months old) cultures consisting of dehydrated hyphae and dormant conidia. Fresh (<3 weeks old) biomasses were toxic more than 10 times towards mammalian cell lines (PK-15 and MNA) compared to the old dormant, dry biomasses, when calculated per biomass wet weight and per conidial particle. Surfactant activity was emitted in exudates from fresh biomass of T. atroviride, Rhizopus sp. and Stachybotrys sp. Surfactant activity was also provoked by fresh conidia from T. atroviride and Stachybotrys sp. strains. Water repealing substances were emitted by cultures of P. expansum, T. atroviride and C. globosum strains. The metabolic state of the indoor fungal growth may influence emission of liquid soluble bioreactive metabolites into the indoor air.

Although transpiration-driven transport of xylem sap is well known to operate under absolute negative pressure, many terrestrial, vascular plants show positive xylem pressure above atmospheric pressure on a seasonal or daily basis, or during early developmental stages. The actual location and mechanisms behind positive xylem pressure remain largely unknown, both in plants that show seasonal xylem pressure before leaf flushing, and those that show a diurnal periodicity of bleeding and guttation. Available evidence shows that positive xylem pressure can be driven based on purely physical forces, osmotic exudation into xylem conduits, or hydraulic pressure in parenchyma cells associated with conduits. The latter two mechanisms may not be mutually exclusive and can be understood based on a similar modelling scenario. Given the renewed interest in positive xylem pressure, this review aims to provide a constructive way forward by discussing similarities and differences of mechanistic models, evaluating available evidence for hydraulic functions, such as rehydration of tissues, refilling of water stores, and embolism repair under positive pressure, and providing recommendations for future research, including methods that avoid or minimise cutting artefacts.