Functional Plant Biology Functional Plant Biology Society
Plant function and evolutionary biology

Functional Plant Biology

Functional Plant Biology

Functional Plant Biology publishes new and significant information on the functional biology of plants at all scales from the molecular through whole plant to community. Read more about the journalMore

Editor-in-Chief: Sergey Shabala

Current Issue

Functional Plant Biology

Volume 44 Number 5 2017

FP16422Why do plants lack sodium pumps and would they benefit from having one?

Jesper T. Pedersen and Michael Palmgren
pp. 473-479

In contrast to their ancestors, vascular plants do not have a Na+ pump. The lack of a Na+ pump might give a hint as to why vascular plants are so sensitive to salinity. In this minireview, we discuss the feasibility of introducing a Na+ pump to increase the salt tolerance of vascular plants.


The accumulation of free polyamines (PAs), the terminal oxidation of PAs and the production of H2O2 by diamine oxidases (DAOs) and polyamine oxidases (PAOs), and the expression of selective DAO and PAO genes, depends on the intensity of salt stress in tomato. H2O2 generated by DAO and PAO in cell walls under lethal salt stress induces cell death caused by irreversible loss of electrolytes from tissues. MDL-72527, a PAO inhibitor, impeded PAO-generated H2O2 production and H2O2-induced nitric oxide accumulation but increased electrolyte leakage from tissues, and thus does not increase salt tolerance.

FP16368Effect of FLOWERING LOCUS C on seed germination depends on dormancy

Logan Blair, Gabriela Auge and Kathleen Donohue
pp. 493-506

The timing of seed germination and flowering influences the fitness of plants in seasonal environments. Two major flowering-time genes – FLOWERING LOCUS C and FRIGIDA – were shown to influence the propensity of seeds to germinate, both immediately after dispersal and after dormancy breakage and re-induction; these genetic differences in dormancy were manifest as genetic differences in temperature-dependent germination. Therefore, flowering and germination share some genetic basis, potentially coordinating or constraining how plants respond to seasonal cues across their life cycle.


It is now accepted that there is continuous carbohydrate unloading and reloading along the entire phloem pathway linking sources and sinks. In what way this affects solute concentration and hydrostatic pressure along the pathway is unknown, and at present this is inaccessible to measurement. Hence modelling of this flow is the only route available. In this paper we use a detailed mechanistic model, able to incorporate lateral flows, to determine the effect of phloem pathway unloading/reloading. With adequate reloading, our calculations indicate that this has no overall effect on equilibrium flows, but has a large effect if unloading is not matched by adequate reloading.

FP15381Calcium improves apoplastic–cytosolic ion homeostasis in salt-stressed Vicia faba leaves

Sherif H. Morgan, Sylvia Lindberg, Pooja Jha Maity, Christoph-Martin Geilfus, Christoph Plieth and Karl-Hermann Mühling
pp. 515-524

This paper describes a new method to measure free Ca concentrations in plants’ extracellular matrix (the apoplast). Extra Ca supply during cultivation increases Ca concentration both in the apoplast and intracellularly, and acidifies the apoplastic pH, which improves ion homeostasis. This may be a further explanation for the favourable effect of Ca for cell expansion and plant growth under salinity.

FP16262Overgrowth (Della) mutants of wheat: development, growth and yield of intragenic suppressors of the Rht-B1c dwarfing gene

Adinda P. Derkx, Carol A. Harding, Asemeh Miraghazadeh and Peter M. Chandler
pp. 525-537

The increased wheat yields that occurred during the Green Revolution were made possible by incorporating semidwarfing alleles of the wheat Della gene into new wheat varieties. These alleles are still in widespread use in current wheat varieties, but we have now isolated many new mutants of this gene and characterised their effects on growth, grain dormancy and yield. The results provide insight into regulation of growth by the DELLA protein and indicate particular alleles of potential value in wheat breeding.


Pre-anthesis water-deficit stress causes detrimental effects on the production of crops such as durum wheat in rain-fed areas. In stress tolerant varieties, the regulation of microRNA160 and the mRNA that it targets, auxin response factors, are potentially associated with the unaffected leaf relative water content and chlorophyll content, and the co-ordinated control of stomatal aperture, which ultimately contribute to the maintenance of grain number and yield. Together, these findings suggest the importance of durum microRNA regulatory modules in water stress responses and provide useful information for improving stress tolerance in breeding.


Annual pasture legumes with a superior ability to acquire soil phosphorus (P) through specialised root systems could lower P-fertiliser requirements. Root traits related to P-acquisition are commonly examined for plants in pots grown in glasshouses; however, root length was greatly decreased and root exudates greatly increased when shoots of these plants were constrained from spreading laterally, as would also occur in dense pasture swards in the field. We suggest that canopy constraint should be routinely used when screening plants in the glasshouse for root traits likely to improve P-acquisition under field conditions.

Online Early

The peer-reviewed and edited version of record published online before inclusion in an issue

Published online 26 April 2017

FP16395Flood tolerance of wheat – the importance of leaf gas films during complete submergence

Anders Winkel, Max Herzog, Dennis Konnerup, Anja Heidi Floytrup and Ole Pedersen
 

Climate changes result in more floods also in regions where the mean precipitation is predicted to decline. Dryland crops such as wheat and barley are particularly vulnerable to submergence stress. Here, we show that superhydrophobic leaf surfaces enhance survival of completely submerged wheat by formation of thin leaf gas films that helps wheat to ‘breathe’ under water.


Methoxypyrazines are responsible for some herbaceous characters in wines of some grape varieties characters and this is determined by the differential expression of a methyltransferase gene known as VvOMT3. In order to investigate the chromatin arrangement of the VvOMT3 gene, histone modifications were studied in the locus and these differed spatially between the skin and flesh tissues, temporally during fruit development and also amongst different VvOMT3 alleles. This study provides evidence of histone tail modification of the VvOMT3 locus in grapevine, which may play a role in the varietal, spatial and developmental regulation of the expression of this gene.

Published online 19 April 2017

FP16339Electrical signalling in Nitellopsis obtusa: potential biomarkers of biologically active compounds

Vilma Kisnieriene, Indre Lapeikaite and Vilmantas Pupkis
 

The electrophysiological response pattern of Nitellopsis obtusa cell can be assessed to evaluate the effect of many biologically active compounds. We illustrate a variety of electrophysiological approaches for the investigations of electrical signaling after chemical treatment in vivo. The insights about the Characean model system are likely to hold for plants in general and even deepen the understanding of the plant evolution.

Published online 18 April 2017

FP16314Endosperm-specific OsPYL8 and OsPYL9 act as positive regulators of the ABA signaling pathway in rice seed germination

Ziqiang Chen, Lan Kong, Yun Zhou, Zaijie Chen, Dagang Tian, Yan Lin, Feng Wang and Songbiao Chen
 

Pyrabactin resistance-like (PYL) proteins were identified as receptors of the plant hormone ABA, but the functions of many PYLs remains to be elucidated. In this work, we determined that rice OsPYL8 and OsPYL9 are specifically expressed in the endosperms. Analyses of transgenic rice plants overexpressing OsPYL8 or OsPYL9 further showed that the two OsPYLs function as positive regulators of ABA signaling pathway in rice seed germination.

Published online 13 April 2017

FP16293Proton and anion transport across the tonoplast vesicles in bromeliad species

Paula Natália Pereira, James Andrew Charles Smith, Eduardo Purgatto and Helenice Mercier
 

Although Crassulacean acid metabolism (CAM) has been well studied in the family Bromeliaceae, the relationship between CAM activity and vacuolar organic-acid transport at night has not. We investigated ATP- and PPi-dependent proton transport in the tonoplast membrane vesicles of seven bromeliad species. We found that tonoplast ATP-driven H+ activity is greater than tonoplast PPi. Our results demonstrate a close correlation between CAM rhythm and the intrinsic properties of vacuolar membranes.

Published online 13 April 2017

FP16324Molecular insights into the functional role of nitric oxide (NO) as a signal for plant responses in chickpea

Parankusam Santisree, Pooja Bhatnagar-Mathur and Kiran K. Sharma
 

Although many studies established nitric oxide (NO) as a signaling molecule in plants, the identification of target molecules of NO has remained elusive due to the lack of in depth molecular studies. Our quantitative proteome analysis suggests the differential regulation of 248 proteins and dynamic regulation of metabolic pathways by exogenous NO donor in chickpea. This is the first report in legumes pointing at the potential candidates that attribute the reported functions of NO in plants.

Published online 10 April 2017

FP16365Morpho-physiological responses to dehydration stress of perennial ryegrass and tall fescue genotypes

Reihaneh Shahidi, Junko Yoshida, Mathias Cougnon, Dirk Reheul and Marie-Christine Van Labeke
 

Climate change will shift the adaptive regions of most forage grasses. We investigated the responses of two tall fescue genotypes and one perennial ryegrass to increasing drought and found that at the physiological and metabolic levels drought was imposed more slowly in tall fescue. Tall fescue may fit into future demands for forage grasses due to its good drought resistance.

Published online 05 April 2017

FP16300Evaluation and application of a targeted SPE-LC-MS method for quantifying plant hormones and phenolics in Arabidopsis

Florence Guérard, Linda de Bont, Bertrand Gakière and Guillaume Tcherkez
 

The quantitation of plant hormones by LC-MS remains challenging because they belong to different chemical classes with contrasted analytical imperatives, and matrix effects often compromise reliable measurements. Here we present a technique based on accurate-mass LC-MS taking advantage of matrix elimination by solid phase extraction before analysis. This method is applied to Arabidopsis samples and as expected, shows that abscisic acid is accumulated under drought and that mutants affected in aspartate oxidase have an increased content in salicylate.

Published online 03 April 2017

FP16164Use of multicolour fluorescence imaging for diagnosis of bacterial and fungal infection on zucchini by implementing machine learning

Mónica Pineda, María Luisa Pérez-Bueno, Vanessa Paredes and Matilde Barón
 

Remote sensing has become an essential tool in phenotyping and precision agriculture to minimise the impact of cultural management on environment and human health. Several imaging techniques are currently in use for the detection of plant stress, but the information generated requires of increasingly advanced mathematical tools. Multicolour fluorescence imaging appears as a promising tool as data provider to feed mathematical models classifying healthy and infected zucchini plants.

Published online 03 April 2017

FP16362Photosynthetic acclimation to elevated CO2 concentration in a sweet pepper (Capsicum annuum) crop under Mediterranean greenhouse conditions: influence of the nitrogen source and salinity

Manuel E. Porras, Pilar Lorenzo, Evangelina Medrano, María J. Sánchez-González, Ginés Otálora-Alcón, María C. Piñero, Francisco M. del Amor and M. Cruz Sánchez-Guerrero
 

An increased CO2 concentration in the atmosphere improves yield of the horticultural crops but its efficiency is reduced after long-term exposure. We studied the evolution of the sweet pepper crop response to CO2 enrichment in Mediterranean greenhouse conditions under different salinity levels. The study revealed the crop phase when high CO2 concentration acclimation occurs and, thus, the point when removal of the CO2 supply could improve its efficiency.

Published online 31 March 2017

FP16048Annual patterns of xylem embolism in high-yield apple cultivars

Barbara Beikircher and Stefan Mayr
 

Winter and spring are crucial periods for long-lived woody species of temperate regions because of the considerable impact on the plants’ water relations. In this study on high-yield apple cultivars, we show which climatic and species-specific factors are primarily responsible for the impairment of the water transport system in winter as well as its restoration in spring. Our findings contribute to the general knowledge about tree hydraulics but are also of relevance for fruit growers.

Published online 29 March 2017

FP16327Anatomical and biochemical characterisation of a barrier to radial O2 loss in adventitious roots of two contrasting Hordeum marinum accessions

Lukasz Kotula, Lukas Schreiber, Timothy D. Colmer and Mikio Nakazono
 

A barrier to radial oxygen loss (ROL) in roots is an important adaptation of many wetland plants growing in waterlogged, anoxic soils; however, knowledge of the nature of the barrier is sparse. The ROL barrier enhances longitudinal oxygen diffusion through aerenchyma to the root tip. Our comparison of two Hordeum marinum accessions differing in ROL barrier strength showed that the deposition of suberin into walls of the root exodermis is associated with reduction in loss of oxygen from basal root zones to the external medium.

Published online 29 March 2017

FP16379Photochemical activity changes accompanying the embryogenesis of pea (Pisum sativum) with yellow and green cotyledons

Galina Smolikova, Vladimir Kreslavski, Olga Shiroglazova, Tatiana Bilova, Elena Sharova, Andrej Frolov and Sergei Medvedev
 

We studied the dynamics of photochemical activity in seed coats and cotyledons during development of yellow- and green-seeded pea cultivars by using the pulse amplitude modulation fluorometric analysis. The fast transients of the chlorophyll a fluorescence revealed higher photochemical activity in the coats of yellow-seeded cultivar at the early- and middle cotyledon stages of seed development in comparison to those observed in the green-seeded ones. Photochemical activity in the cotyledons of both cultivars could not be any more detected at the late cotyledon stage. This process was triggered by dehydration of seed tissues.

Published online 29 March 2017

FP16357Environmental factors constraining adventitious root formation during flooding of Solanum dulcamara

Qian Zhang, Heidrun Huber, Jannah W. T. Boerakker, Daniek Bosch, Hans de Kroon and Eric J. W. Visser
 

Flooding generally poses a threat to terrestrial plants, but wetland species display adaptations that prevent damage by the adverse conditions imposed by floods. One of these adaptations – formation of adventitious roots that replace the original non-adapted root system – may be severely constrained if insufficient light or contact of leaves with the atmosphere is present during the flooding event. This results in significantly fewer adventitious roots growing out of the stem, and thus poor performance in these stressful conditions.


Root growth is controlled by phytohormones, but what cellular processes are regulated and how it occurs is still an open question. Here it is shown that cytokinin affects root growth mainly through its effect on cell proliferation, and does not initiate the transition of cells to differentiation as previously thought. Cellular analysis performed could be applied for the analysis of how any plant hormone influences developmental processes in plant roots.

Published online 24 March 2017

FP16283Cyclosis-mediated long distance communications of chloroplasts in giant cells of Characeae

Anna V. Komarova, Vladimir S. Sukhov and Alexander A. Bulychev
 

Intracellular communications in plant cells of large dimensions rely primarily on cytoplasmic streaming, because diffusion is too slow for the transport on mm-scale distances. Illumination of a small cell spot at a various distances from the point of chlorophyll fluorescence measurements revealed the wave-like propagation of the fluorescence response along the cell length. The results show that the photosynthetic function of immobile chloroplasts under constant light can be affected by long-distance transmission of a photosynthetically active metabolite from the remote cell parts.

Published online 23 March 2017

FP16401Leaf gas film retention during submergence of 14 cultivars of wheat (Triticum aestivum)

Dennis Konnerup, Anders Winkel, Max Herzog and Ole Pedersen
 

Some terrestrial plants, including wheat (Triticum aestivum), possess superhydrophobic leaf surfaces that retain a thin gas film when submerged. We tested gas film retention time of 14 different wheat cultivars and found that wheat could retain the gas films for a minimum of 2 days. We suggest that leaf gas film is a relevant trait to use as a selection criterion to improve the flood tolerance of crops that become temporarily submerged.


Understanding the molecular mechanisms of plant development constitutes an important field of investigations in the current era of plant biology research. Nitric oxide signalling regulates a variety of biochemical processes in plants. Present review provides an in-depth analysis of our current understanding on the subject, particularly with reference to plant growth under stress conditions.

Published online 22 March 2017

FP16384Melatonin in plant signalling and behaviour

Lauren A. E. Erland, Praveen K. Saxena and Susan J. Murch
 

Melatonin is an important hormone and signalling molecule in all forms of life including humans, plants and bacteria. Recent plant physiology and genomic experiments have described the redirection of plant growth and metabolism, and demonstrated a diversity of genes involved in response to melatonin, however, the exact metabolic cascades that translate melatonin signals into physiological responses is not fully understood. This review provides an overview of melatonin mediated signalling manifested as behaviours and its roles in basic and industrial research.

Published online 16 March 2017

FP16326Sunpatiens compact hot coral: memristors in flowers

Alexander G. Volkov and Eunice K. Nyasani
 

Memristors, or resistors with memory, exist in vivo as components of plasma membranes in plants, fruits, roots and seeds. Authors found memristors in an androecium, spur, petals and pedicel in Sunpatiens flowers. The discovery of memristors in Sunpatiens (Impatiens spp.) creates a new direction in the modelling and understanding of electrophysiological phenomena and memory elements in flowers.

Published online 14 March 2017

FP16348Phloem fibres as motors of gravitropic behaviour of flax plants: level of transcriptome

Oleg Gorshkov, Natalia Mokshina, Nadezda Ibragimova, Marina Ageeva, Natalia Gogoleva and Tatyana Gorshkova
 

Plant fibres with a tertiary cell wall (G-layer) may function as plant ‘muscles’. Large-scale transcriptome profiling of isolated flax phloem fibres permitted to identify the major players and regulatory elements that operate during graviresponce specifically in the fibres of the pulling stem side. The suggested mechanisms of phloem fibre involvement in tropisms may considerably renew the concept of herbaceous plant behaviour upon gravistimulation.

Published online 14 March 2017

FP16318Plant ion channels and transporters in herbivory-induced signalling

Shuitian Luo, Xiao Zhang, Jinfei Wang, Chunyang Jiao, Yingying Chen and Yingbai Shen
 

Clarifying herbivory-induced plant cellular signalling is a critical step to push the research of plant-herbivore interaction forward. We review the role of ion channels/transporters in modulating herbivory-induced early signalling events and rapid systemic signal transmission in plants. This work provides a comprehensive source of information about plant defensive strategies upon attack.

Published online 14 March 2017

FP16292Studies of cytokinin receptor–phosphotransmitter interaction provide evidences for the initiation of cytokinin signalling in the endoplasmic reticulum

Sergey N. Lomin, Yulia A. Myakushina, Dmitry V. Arkhipov, Olga G. Leonova, Vladimir I. Popenko, Thomas Schmülling and Georgy A. Romanov
 

Cytokinin is an important plant hormone and its mode of action has been extensively studied; however, to date, the subcellular localisation of cytokinin perception and signal transduction remains a matter of debate. This study describes cytokinin receptor–phosphotransmitter interaction and its subcellular localisation in living plant cells and it provides several experimental evidences for receptor activity at the endoplasmic reticulum (ER) membrane. It is concluded that intracellular cytokinins within the ER lumen may play an important role in cytokinin signalling, at least in some cell types.

Published online 07 March 2017

FP16377Cell differentiation in nitrogen-fixing nodules hosting symbiosomes

Anna V. Tsyganova, Anna B. Kitaeva and Viktor E. Tsyganov
 

Rhizobium bacteria, which live within the root nodules of legumes, allow plants to capture nitrogen gas from the atmosphere and use it for their own growth. Central to this symbiosis is an intracellular structure, called the symbiosome, in which nitrogen-fixing bacterial cells exchange components with the host cells that harbor them. Recent research on the differentiation of symbiosomes and of the infected cells that accommodate them has helped to decipher some general molecular mechanisms of cell differentiation.

Published online 03 March 2017

FP16342Parameters of electrical signals and photosynthetic responses induced by them in pea seedlings depend on the nature of stimulus

Vladimir Vodeneev, Maxim Mudrilov, Elena Akinchits, Irina Balalaeva and Vladimir Sukhov
 

Plants, like animals, produce electrical signals in response to various external influences. In this study we raised a question whether the electrical signals transmit information about the nature of the stimulus, and found out that different stimuli induce signals of varied parameters. The obtained results explain how plants adapt to changing environment.

Published online 28 February 2017

FP16380The role of ion disequilibrium in induction of root cell death and autophagy by environmental stresses

Vadim Demidchik, Elena V. Tyutereva and Olga V. Voitsekhovskaja
 

Environmental stresses are main causes for low agricultural productivity. At the cellular level, stresses induce generation of reactive oxygen species (ROS), ion disequilibrium, autophagy and programmed cell death (PCD). Here we propose that these processes interact and that ROS and ion disequilibrium are triggers of autophagy and PCD. Overall, presented data contribute to understanding plant stress physiology.


Transient elevation of cytosolic Ca2+, also referred as a Ca2+signal, is as central phenomenon of plant signalling. Plants evolved sophisticated systems to initiate, amplify and terminate Ca2+ signals. Structure and properties of these systems, including Ca2+-permeable ion channels, Ca2+-ATPases, Ca2+/H+ exchangers and ‘ROS-Ca2+ hub’ are discussed here. They provide a fine-tuned mechanism for encoding diverse external and internal stimuli.

Published online 13 February 2017

FP16347Spatial distribution of organelles in leaf cells and soybean root nodules revealed by focused ion beam-scanning electron microscopy

Brandon C. Reagan, Paul J. -Y. Kim, Preston D. Perry, John R. Dunlap and Tessa M. Burch-Smith
 

Focussed ion bean scanning electron microscopy (FIB-SEM) is a technique that can be used to generate 3D renderings of cells and their contents. Although FIB-SEM has been regularly used to investigate animal cells and tissues, it has rarely been deployed to study plant structures. Here we demonstrate that FIB-SEM can easily be used to study plant samples and have discovered previously unknown arrangements of organelles and membranes in those samples.

Published online 03 February 2017

FP16322Formation mechanisms of superoxide radical and hydrogen peroxide in chloroplasts, and factors determining the signalling by hydrogen peroxide

Boris N. Ivanov, Maria M. Borisova-Mubarakshina and Marina A. Kozuleva
 

Photosynthetic electron transport chain is not only source of ATP and NADPH for photosynthesis; it is a sensor, informing adaptation systems of plant about environmental changes. An important transmitter of this information is hydrogen peroxide whose mechanisms of formation are presented, laying special emphasis on the formation outside and within thylakoid membrane. It is discussed, that the formation place can ensure definite signal about the specific environmental change.

Published online 11 January 2017

FP16321Arabidopsis thaliana phytaspase: identification and peculiar properties

Nina V. Chichkova, Raisa A. Galiullina, Larisa V. Mochalova, Svetlana V. Trusova, Zulfazli M. Sobri, Patrick Gallois and Andrey B. Vartapetian
 

Although plant proteases of the phytaspase family are important contributors to stress-induced plant cell death, phytaspase of a classical model plant Arabidopsis thaliana has escaped identification thus far. We identified the Arabidopsis phytaspase-encoding gene and characterised the recombinant enzyme. Substrate specificity and properties of the Arabidopsis phytaspase display both important similarities with and distinctions from the already characterised phytaspases.

Published online 20 December 2016

FP16337cGMP signalling in plants: from enigma to main stream

Jean-Charles Isner and Frans J. M. Maathuis
 

Cyclic GMP (cGMP) signalling in plants is crucial for many physiological processes. Recent analytical and genomic developments now allow detailed studies into the biochemistry and physiological role of cGMP in plants, and the latest findings are reviewed in this article.

Published online 16 December 2016

FP16338Two-pore cation (TPC) channel: not a shorthanded one

Igor Pottosin and Oxana Dobrovinskaya
 

Large conductance SV/TPC1 channels are ubiquitously and abundantly expressed in the vacuolar membranes of higher plants. They are unique established Ca2+-permeable channels in vacuoles, but their activity is strongly negatively controlled, so that they were believed to be inactive or to act only locally. Recent evidence suggests the key role of SV/TPC1 channels in the long-distance Ca2+ signalling.


Plants adapt to environmental light conditions with the use of the sophisticated phytochrome system. In this work, polymorphism of its major component – phytochrome A– was investigated. With the use of transgenic Arabidopsis and fluorescence technique, it was shown that two molecular types of the photoreceptor differ by the state of phosphorylation and their existence accounts for its complex functions.

Published online 09 November 2016

FP16242Rapid changes in root HvPIP2;2 aquaporins abundance and ABA concentration are required to enhance root hydraulic conductivity and maintain leaf water potential in response to increased evaporative demand

Dmitry S. Veselov, Dmitry S. Veselov, Guzel V. Sharipova, Guzel V. Sharipova, Stanislav Yu. Veselov, Stanislav Yu. Veselov, Ian C. Dodd, Ian C. Dodd, Igor Ivanov, Igor Ivanov, Guzel R. Kudoyarova and Guzel R. Kudoyarova
 

The ABA-deficient barley mutant Az34 and wild type (WT) were exposed to air warming. Although transpiration rate of both genotypes increased, leaf water potential decreased in the mutant but was maintained in WT plants. Only WT plants showed increased root ABA accumulation, which increased root hydraulic conductivity and aquaporin abundance, which seems important in maintaining leaf hydration.

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