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Plants can activate protective and defense mechanisms under biotic and abiotic stresses. Their roots naturally grow in the soil, but when they encounter sunlight in the top-soil layers, they may move away from the light source to seek darkness. Here we investigate the skototropic behavior of roots, which promotes their fitness and survival. Glutamate-like receptors (GLRs) of plants play roles in sensing and responding to signals, but their role in root skototropism is not yet understood. Light-induced tropisms are known to be affected by auxin distribution, mainly determined by auxin efflux proteins (PIN proteins) at the root tip. However, the role of PIN proteins in root skototropism has not been investigated yet. To better understand root skototropism and its connection to the distance between roots and light, we established five distance settings between seedlings and darkness to investigate the variations in root bending tendencies. We compared differences in root skototropic behavior across different expression lines of Arabidopsis thaliana seedlings (atglr3.7 ko, AtGLR3.7 OE, and pin2 knockout) to comprehend their functions. Our research shows that as the distance between roots and darkness increases, the root’s positive skototropism noticeably weakens. Our findings highlight the involvement of GLR3.7 and PIN2 in root skototropism.

Abstract Background Shifts in dynamic equilibria of the abundance of cellular molecules in plant-pathogen interactions need further exploration. We induced PTI in optimally growing Arabidopsis thaliana seedlings for 16 h, returning them to growth conditions for another 16 h. Methods Turn-over and abundance of 99 flg22 responding proteins were measured chronologically using a stable heavy nitrogen isotope partial labeling strategy and targeted liquid chromatography coupled to mass spectrometry (PRM LC–MS). These experiments were complemented by measurements of mRNA and phytohormone levels. Results Changes in synthesis and degradation rate constants (K s and K d) regulated tryptophane and glucosinolate, IAA transport, and photosynthesis-associated protein (PAP) homeostasis in growth/PTI transitions independently of mRNA levels. K s values increased after elicitation while protein and mRNA levels became uncorrelated. mRNA returned to pre-elicitation levels, yet protein abundance remained at PTI levels even 16 h after media exchange, indicating protein levels were robust and unresponsive to transition back to growth. The abundance of 23 PAPs including FERREDOXIN-NADP( +)-OXIDOREDUCTASE (FNR1) decreased 16 h after PAMP exposure, their depletion was nearly abolished in the myc234 mutant. FNR1 K d increased as mRNA levels decreased early in PTI, its K s decreased in prolonged PTI. FNR1 K d was lower in myc234, mRNA levels decreased as in wild type. Conclusions Protein K d and K s values change in response to flg22 exposure and constitute an additional layer of protein abundance regulation in growth defense transitions next to changes in mRNA levels. Our results suggest photosystem remodeling in PTI to direct electron flow away from the photosynthetic carbon reaction towards ROS production as an active defense mechanism controlled post-transcriptionally and by MYC2 and homologs. Target proteins accumulated later and PAP and auxin/IAA depletion was repressed in myc234 indicating a positive effect of the transcription factors in the establishment of PTI.

Diverse branching forms have evolved multiple times across the tree of life to facilitate resource acquisition and exchange with the environment. In the vascular plant group, the ancestral pattern of branching involves dichotomy of a parent shoot apex to form two new daughter apices. The molecular basis of axillary branching in Arabidopsis is well understood, but few regulators of dichotomous branching are known. Through analyses of dichotomous branching in the lycophyte, Selaginella kraussiana, we identify PIN-mediated auxin transport as an ancestral branch regulator of vascular plants. We show that short-range auxin transport out of the apices promotes dichotomy and that branch dominance is globally coordinated by long-range auxin transport. Uniquely in Selaginella, angle meristems initiate at each dichotomy, and these can develop into rhizophores or branching angle shoots. We show that long-range auxin transport and a transitory drop in PIN expression are involved in angle shoot development. We conclude that PIN-mediated auxin transport is an ancestral mechanism for vascular plant branching that was independently recruited into Selaginella angle shoot development and seed plant axillary branching during evolution. [ABSTRACT FROM AUTHOR]

Auxin, which is transported in the outermost cell layer, is one of the major players involved in plant organ initiation and positioning at the shoot apical meristem (SAM). However, recent studies have recognized the role of putative internal signals as an important factor collaborating with the well-described superficial pathway of organogenesis regulation. Different internal signals have been proposed; however, their nature and transport route have not been precisely determined. Therefore, in this mini-review, we aimed to summarize the current knowledge regarding the auxin-dependent regulation of organ positioning at the SAM and to discuss the vascular system as a potential route for internal signals. In addition, as regular organ patterning is a universal phenomenon, we focus on the role of the vasculature in this process in the major lineages of land plants, i.e., bryophytes, lycophytes, ferns, gymnosperms, and angiosperms. [ABSTRACT FROM AUTHOR]

Summary: Photomorphogenic remodelling of seedling growth is a key developmental transition in the plant life cycle. The α/β‐hydrolase signalling protein KARRIKIN‐INSENSITIVE2 (KAI2), a close homologue of the strigolactone receptor DWARF14 (D14), is involved in this process, but it is unclear how the effects of KAI2 on development are mediated.Here, using a combination of physiological, pharmacological, genetic and imaging approaches in Arabidopsis thaliana (Heynh.) we show that kai2 phenotypes arise because of a failure to downregulate auxin transport from the seedling shoot apex towards the root system, rather than a failure to respond to light per se.We demonstrate that KAI2 controls the light‐induced remodelling of the PIN‐mediated auxin transport system in seedlings, promoting a reduction in PIN7 abundance in older tissues, and an increase of PIN1/PIN2 abundance in the root meristem. We show that removing PIN3, PIN4 and PIN7 from kai2 mutants, or pharmacological inhibition of auxin transport and synthesis, is sufficient to suppress most kai2 seedling phenotypes.We conclude that KAI2 regulates seedling morphogenesis by its effects on the auxin transport system. We propose that KAI2 is not required for the light‐mediated changes in PIN gene expression but is required for the appropriate changes in PIN protein abundance within cells. [ABSTRACT FROM AUTHOR]

Pelargonic acid (PA) is a saturated fatty acid commonly found in several organisms, that is known for its phytotoxic effect and its use as bioherbicide for sustainable weed management. Although PA is already commercialised as bioherbicide, its molecular targets and mode of action is unknown according to the Herbicide Resistance Action Committee. Therefore, the aim of this work was focusing on the way this natural active substance impacts the plant metabolism of the model species Arabidopsis thaliana. PA caused increase of secondary and adventitious roots, as well as torsion, loss of gravitropism and phytotoxic effects. Moreover, PA altered the cellular arrangement and the PIN proteins activity. Computational simulations revealed that the intermolecular interactions between PA and the polar auxin transporter protein PIN1 are very similar to those established between the natural auxin IAA and PIN1. However, under intracellular conditions, the PA-PIN1 binding is more energetically stable than the IAA-PIN1. These results suggest that PA could act as an auxin-mimics bioherbicide. The exogenous application of PA would be responsible for the alterations observed both at structural and ultrastructural levels, which would be caused by the alteration on the transport of auxins into the plant, inducing root inhibition and ultimately total stop of root growth. • Pelargonic acid is a natural metabolite that can alter root architecture. • This specialised metabolite causes alterations in auxin distribution. • Pelargonic acid shares a binding site with IAA on the PIN1 protein. • The results suggest that PA could act as a bioherbicide by altering the auxin polar transport. [ABSTRACT FROM AUTHOR]

Plants can activate protective and defense mechanisms under biotic and abiotic stresses. Their roots naturally grow in the soil, but when they encounter sunlight in the top-soil layers, they may move away from the light source to seek darkness. Here we investigate the skototropic behavior of roots, which promotes their fitness and survival. Glutamate-like receptors (GLRs) of plants play roles in sensing and responding to signals, but their role in root skototropism is not yet understood. Light-induced tropisms are known to be affected by auxin distribution, mainly determined by auxin efflux proteins (PIN proteins) at the root tip. However, the role of PIN proteins in root skototropism has not been investigated yet. To better understand root skototropism and its connection to the distance between roots and light, we established five distance settings between seedlings and darkness to investigate the variations in root bending tendencies. We compared differences in root skototropic behavior across different expression lines of Arabidopsis thaliana seedlings ( atglr3.7 ko, AtGLR3.7 OE , and pin2 knockout ) to comprehend their functions. Our research shows that as the distance between roots and darkness increases, the root's positive skototropism noticeably weakens. Our findings highlight the involvement of GLR3.7 and PIN2 in root skototropism.

Summary: The plant hormone auxin and its directional intercellular transport play a major role in diverse aspects of plant growth and development. The establishment of auxin gradients requires the asymmetric distribution of members of the auxin efflux carrier PIN‐FORMED (PIN) protein family to the plasma membrane. An endocytic pathway regulates the recycling of PIN proteins between the plasma membrane and endosomes, providing a mechanism for dynamic localisation.N‐Ethylmaleimide‐sensitive factor adaptor protein receptors (SNAP receptors, SNAREs) mediate fusion between vesicles and target membranes and are classed as Q‐ or R‐SNAREs based on their sequence. We analysed gain‐ and loss‐of‐function mutants, dominant‐negative transgenics and localisation of the Arabidopsis R‐SNARE VAMP714 protein to understand its function.We demonstrate that VAMP714 is essential for the insertion of PINs into the plasma membrane, for polar auxin transport, root gravitropism and morphogenesis. VAMP714 gene expression is upregulated by auxin, and the VAMP714 protein co‐localises with endoplasmic reticulum and Golgi vesicles and with PIN proteins at the plasma membrane.It is proposed that VAMP714 mediates the delivery of PIN‐carrying vesicles to the plasma membrane, and that this forms part of a positive regulatory loop in which auxin activates a VAMP714‐dependent PIN/auxin transport system to control development. [ABSTRACT FROM AUTHOR]

In epiphytes, aerial roots are important to combat water-deficient, nutrient-poor, and high-irradiance microhabitats. However, whether aerial roots can respond to gravity and whether auxin plays a role in regulating aerial root development remain open-ended questions. Here, we investigated the gravitropic response of the epiphytic orchid Phalaenopsis aphrodite. Our data showed that aerial roots of P. aphrodite failed to respond to gravity, and this was correlated with a lack of starch granules/statolith sedimentation in the roots and the absence of the auxin efflux carrier PIN2 gene. Using an established auxin reporter, we discovered that auxin maximum was absent in the quiescent center of aerial roots of P. aphrodite. Also, gravity failed to trigger auxin redistribution in the root caps. Hence, loss of gravity sensing and gravity-dependent auxin redistribution may be the genetic factors contributing to aerial root development. Moreover, the architectural and functional innovations that achieve fast gravitropism in the flowering plants appear to be lost in both terrestrial and epiphytic orchids, but are present in the early diverged orchid subfamilies. Taken together, our findings provide physiological and molecular evidence to support the notion that epiphytic orchids lack gravitropism and suggest diverse geotropic responses in the orchid family., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Biostimulants may be particularly interesting for application in agricultural and horticultural crops since they can exert a growth-promoting effect on roots. This may be important for promoting longitudinal and lateral root growth and therefore increasing belowground vegetative growth, which may in turn lead to improved aboveground vegetative growth and increased yields. Here, we examined the effects and mechanism of action of an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) on the root growth of tomato plants, with an emphasis on its possible role on chorismate-derived hormones (auxin, salicylic acid, and melatonin). Tomato plants growing in hydroponic systems were exposed to either nutrient stress conditions (experiment 1) or suboptimal temperatures (experiment 2) in a greenhouse, and the concentration of auxin, salicylic acid, and melatonin in roots were measured just prior and after the application of the biostimulant. Results showed that the application of Pepton exerted a growth-promoting effect on roots in plants growing under suboptimal conditions, which might be associated with enhanced salicylic acid levels in roots. The extent of effects of this enzymatically hydrolyzed animal protein-based biostimulant might strongly depend on the growth conditions and stage of root system development. It is concluded that an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) may exert a positive effect enhancing primary and lateral root growth of tomato plants growing under suboptimal conditions, by stimulating the biosynthesis of specific hormonal pathways, such as salicylic acid under stress.

Biostimulants may be particularly interesting for application in agricultural and horticultural crops since they can exert a growth-promoting effect on roots. This may be important for promoting longitudinal and lateral root growth and therefore increasing belowground vegetative growth, which may in turn lead to improved aboveground vegetative growth and increased yields. Here, we examined the effects and mechanism of action of an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) on the root growth of tomato plants, with an emphasis on its possible role on chorismate-derived hormones (auxin, salicylic acid, and melatonin). Tomato plants growing in hydroponic systems were exposed to either nutrient stress conditions (experiment 1) or suboptimal temperatures (experiment 2) in a greenhouse, and the concentration of auxin, salicylic acid, and melatonin in roots were measured just prior and after the application of the biostimulant. Results showed that the application of Pepton exerted a growth-promoting effect on roots in plants growing under suboptimal conditions, which might be associated with enhanced salicylic acid levels in roots. The extent of effects of this enzymatically hydrolyzed animal protein-based biostimulant might strongly depend on the growth conditions and stage of root system development. It is concluded that an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) may exert a positive effect enhancing primary and lateral root growth of tomato plants growing under suboptimal conditions, by stimulating the biosynthesis of specific hormonal pathways, such as salicylic acid under stress. [ABSTRACT FROM AUTHOR]

Norharmane is an indole alkaloid that can be found in several terrestrial plants, as well as in some dinoflagellates and cyanobacteria. The aim of this study was to focus on the way this metabolite impacts the plant metabolism of the model species Arabidopsis thaliana. This metabolite caused increase of secondary and adventitious roots, as well as torsion, toxic effects, and a decrease in root length. Moreover, norharmane altered the cellular arrangement, resulting in unfinished cell walls, decreased auxin content and inhibited PIN proteins activity. All the alterations suggest that norharmane alters polar auxin transport by inhibiting PIN2, PIN3 and PIN7 transport proteins, thus causing a significant inhibitory effect on the growth of A. thaliana seedlings. • Norharmane affects PIN2, PIN3 and PIN7 proteins in Arabidopsis seedlings. • Norharmane inhibition of PIN2, PIN3 and PIN7 proteins results in altered polar auxin transport. • The altered polar auxin transport results in strongly decreased growth. [ABSTRACT FROM AUTHOR]

Figured wood of Karelian birch (Betula pendula Roth var. carelica (Merckl.) Hämet-Ahti) is highly appraised for its ornamental properties. The reasons for its formation remain largely unclear; judging by available data, we are dealing here with auxin inactivation through its interaction with sugars. The aim of this study has been to reveal the correlations between abnormal wood formation in Karelian birch and auxin conjugation producing IAA-glucose. Karelian birch trees with figured and non-figured wood in the trunk were used. Figured plants had a much higher expression of the gene encoding for the enzyme IAA-glucose synthase, which catalyzes IAA-glucose synthesis. The data obtained suggest that auxin conjugation in Karelian birch via a chain of biochemical reactions may be associated with hexoses produced in the apoplast through sucrose cleavage by cell wall invertase. The activity of the enzyme in figured Karelian birch trees is several times higher than in non-figured plants. Vessel differentiation requires free auxin. We assume that a reduced number of vessels in figured wood may be caused by auxin conjugation in the zone of xylem cell growth and differentiation. Figured wood of Karelian birch was found to have an increased transcript level of the PIN3 gene, which encodes for the auxin efflux carrier protein PIN3 responsible for lateral transport of the hormone. In this regard, PIN3 may generate auxin fluxes with a complex configuration. Microscopic analysis provides evidence that PIN3 overexpression in trunk tissues of Karelian birch is associated with active differentiation of parenchyma cells and disruption of wood structure. [ABSTRACT FROM AUTHOR]

Novel molecular pinball machines of the plasma membrane control cytosolic Ca2+ levels that regulate plant metabolism. The essential components involve: 1. an auxin-activated proton pump; 2. arabinogalactan glycoproteins (AGPs); 3. Ca2+ channels; 4. auxin-efflux “PIN” proteins. Typical pinball machines release pinballs that trigger various sound and visual effects. However, in plants, “proton pinballs” eject Ca2+ bound by paired glucuronic acid residues of numerous glycomodules in periplasmic AGP-Ca2+. Freed Ca2+ ions flow down the electrostatic gradient through open Ca2+ channels into the cytosol, thus activating numerous Ca2+-dependent activities. Clearly, cytosolic Ca2+ levels depend on the activity of the proton pump, the state of Ca2+ channels and the size of the periplasmic AGP-Ca2+ capacitor; proton pump activation is a major regulatory focal point tightly controlled by the supply of auxin. Auxin efflux carriers conveniently known as “PIN” proteins (null mutants are pin-shaped) pump auxin from cell to cell. Mechanosensitive Ca2+ channels and their activation by reactive oxygen species (ROS) are yet another factor regulating cytosolic Ca2+. Cell expansion also triggers proton pump/pinball activity by the mechanotransduction of wall stress via Hechtian adhesion, thus forming a Hechtian oscillator that underlies cycles of wall plasticity and oscillatory growth. Finally, the Ca2+ homeostasis of plants depends on cell surface external storage as a source of dynamic Ca2+, unlike the internal ER storage source of animals, where the added regulatory complexities ranging from vitamin D to parathormone contrast with the elegant simplicity of plant life. This paper summarizes a sixty-year Odyssey.

Summary: Gravitropism is an adaptive response that orients plant growth parallel to the gravity vector. Asymmetric distribution of the phytohormone auxin is a necessary prerequisite to the tropic bending both in roots and shoots. During hypocotyl gravitropic response, the PIN3 auxin transporter polarizes within gravity‐sensing cells to redirect intercellular auxin fluxes. First gravity‐induced PIN3 polarization to the bottom cell membranes leads to the auxin accumulation at the lower side of the organ, initiating bending and, later, auxin feedback‐mediated repolarization restores symmetric auxin distribution to terminate bending. Here, we performed a forward genetic screen to identify regulators of both PIN3 polarization events during gravitropic response. We searched for mutants with defective PIN3 polarizations based on easy‐to‐score morphological outputs of decreased or increased gravity‐induced hypocotyl bending. We identified the number of hypocotyl reduced bending (hrb) and hypocotyl hyperbending (hhb) mutants, revealing that reduced bending correlated typically with defective gravity‐induced PIN3 relocation whereas all analyzed hhb mutants showed defects in the second, auxin‐mediated PIN3 relocation. Next‐generation sequencing‐aided mutation mapping identified several candidate genes, including SCARECROW and ACTIN2, revealing roles of endodermis specification and actin cytoskeleton in the respective gravity‐ and auxin‐induced PIN polarization events. The hypocotyl gravitropism screen thus promises to provide novel insights into mechanisms underlying cell polarity and plant adaptive development. Significance Statement: Plant gravitropism involves two distinct cellular polarizations of PIN3 auxin transporter to redirect intercellular auxin fluxes for the bending response. Here we identified a number of Arabidopsis mutants defective specifically in the PIN3 polarization events, establishing that gravity‐induced polarization is required for bending initiation and auxin feedback‐mediated second polarization for its termination. Molecular analysis of selected mutants revealed a crucial role of SCARECROW‐mediated endodermis specification and actin cytoskeleton in these polarization events during gravitropism. [ABSTRACT FROM AUTHOR]

The proper distribution of the hormone auxin is essential for plant development. It is channeled by auxin efflux carriers of the PIN family, typically asymmetrically located on the plasma membrane (PM). Several studies demonstrated that some PIN transporters are also located at the endoplasmic reticulum (ER). From the PM-PINs, they differ in a shorter internal hydrophilic loop, which carries the most important structural features required for their subcellular localization, but their biological role is otherwise relatively poorly known. We discuss how ER-PINs take part in maintaining intracellular auxin homeostasis, possibly by modulating the internal levels of IAA; it seems that the exact identity of the metabolites downstream of ER-PINs is not entirely clear as well. We further review the current knowledge about their predicted structure, evolution and localization. Finally, we also summarize their role in plant development.

Coordination of plant development requires modulation of growth responses that are under control of the phytohormone auxin. PIN-FORMED plasma membrane proteins, involved in intercellular transport of the growth regulator, are key to the transmission of such auxin signals and subject to multilevel surveillance mechanisms, including reversible post-translational modifications. Apart from well-studied PIN protein modifications, namely phosphorylation and ubiquitylation, no further post-translational modifications have been described so far. Here, we focused on root-specific Arabidopsis PIN2 and explored functional implications of two evolutionary conserved cysteines, by a combination of in silico and molecular approaches. PIN2 sequence alignments and modeling predictions indicated that both cysteines are facing the cytoplasm and therefore would be accessible to redox status-controlled modifications. Notably, mutant pin2C-A alleles retained functionality, demonstrated by their ability to almost completely rescue defects of a pin2 null allele, whereas high resolution analysis of pin2C-A localization revealed increased intracellular accumulation, and altered protein distribution within plasma membrane micro-domains. The observed effects of cysteine replacements on root growth and PIN2 localization are consistent with a model in which redox status-dependent cysteine modifications participate in the regulation of PIN2 mobility, thereby fine-tuning polar auxin transport. [ABSTRACT FROM AUTHOR]

Abstract Plant development is exceptionally flexible as manifested by its potential for organogenesis and regeneration, which are processes involving rearrangements of tissue polarities. Fundamental questions concern how individual cells can polarize in a coordinated manner to integrate into the multicellular context. In canalization models, the signaling molecule auxin acts as a polarizing cue, and feedback on the intercellular auxin flow is key for synchronized polarity rearrangements. We provide a novel mechanistic framework for canalization, based on up‐to‐date experimental data and minimal, biologically plausible assumptions. Our model combines the intracellular auxin signaling for expression of PINFORMED (PIN) auxin transporters and the theoretical postulation of extracellular auxin signaling for modulation of PIN subcellular dynamics. Computer simulations faithfully and robustly recapitulated the experimentally observed patterns of tissue polarity and asymmetric auxin distribution during formation and regeneration of vascular systems and during the competitive regulation of shoot branching by apical dominance. Additionally, our model generated new predictions that could be experimentally validated, highlighting a mechanistically conceivable explanation for the PIN polarization and canalization of the auxin flow in plants.

Understanding the mechanisms regulating root development under drought conditions is an important question for plant biology and world agriculture., We examine the effect of osmotic stress on abscisic acid (ABA), cytokinin and ethylene responses and how they mediate auxin transport, distribution and root growth through effects on PIN proteins. We integrate experimental data to construct hormonal crosstalk networks to formulate a systems view of root growth regulation by multiple hormones., Experimental analysis shows: that ABA-dependent and ABA-independent stress responses increase under osmotic stress, but cytokinin responses are only slightly reduced; inhibition of root growth under osmotic stress does not require ethylene signalling, but auxin can rescue root growth and meristem size; osmotic stress modulates auxin transporter levels and localization, reducing root auxin concentrations; PIN1 levels are reduced under stress in an ABA-dependent manner, overriding ethylene effects; and the interplay among ABA, ethylene, cytokinin and auxin is tissue-specific, as evidenced by differential responses of PIN1 and PIN2 to osmotic stress., Combining experimental analysis with network construction reveals that ABA regulates root growth under osmotic stress conditions via an interacting hormonal network with cytokinin, ethylene and auxin. [ABSTRACT FROM AUTHOR]

Coordination of plant development requires modulation of growth responses that are under control of the phytohormone auxin. PIN-FORMED plasma membrane proteins, involved in intercellular transport of the growth regulator, are key to the transmission of such auxin signals and subject to multilevel surveillance mechanisms, including reversible post-translational modifications. Apart from well-studied PIN protein modifications, namely phosphorylation and ubiquitylation, no further post-translational modifications have been described so far. Here, we focused on root-specific Arabidopsis PIN2 and explored functional implications of two evolutionary conserved cysteines, by a combination of in silico and molecular approaches. PIN2 sequence alignments and modeling predictions indicated that both cysteines are facing the cytoplasm and therefore would be accessible to redox status-controlled modifications. Notably, mutant pin2C−A alleles retained functionality, demonstrated by their ability to almost completely rescue defects of a pin2 null allele, whereas high resolution analysis of pin2C−A localization revealed increased intracellular accumulation, and altered protein distribution within plasma membrane micro-domains. The observed effects of cysteine replacements on root growth and PIN2 localization are consistent with a model in which redox status-dependent cysteine modifications participate in the regulation of PIN2 mobility, thereby fine-tuning polar auxin transport.

Compartmentalization of cellular functions relies on partitioning of domains of diverse sizes within the plasma membrane (PM). Macro-domains measure several micrometers and contain specific proteins concentrated to specific sides (apical, basal and lateral) of the PM conferring a polarity to the cell. Cell polarity is one of the driving forces in tissue and growth patterning. To maintain macro-domains within the PM, eukaryotic cells exert diverse mechanisms to counteract the free lateral diffusion of proteins. Protein activation/inactivation, endocytosis, PM recycling of transmembrane proteins and the role of diffusion barriers in macro-domains partitioning at PM will be discussed. Moreover, as plasmodesmata (PDs) are domains inserted within the PM which also mediate tissue and growth patterning, it is essential to understand how segregation of specific set of proteins is maintained at PDs while PDs domains are smaller in size compared to macro-domains. Here, we will present mechanisms allowing restriction of proteins at PM macrodomains, but for which molecular components have been found in PDs proteome. We will explore the hypothesis that partitioning of macro-domains and PDs may be ruled by similar mechanisms.

Although plant and animal cells use a similar core mechanism to deliver proteins to the plasma membrane, their different lifestyle, body organization and specific cell structures resulted in the acquisition of regulatory mechanisms that vary in the two kingdoms. In particular, cell polarity regulators do not seem to be conserved, because genes encoding key components are absent in plant genomes. In plants, the broad knowledge on polarity derives from the study of auxin transporters, the PIN-FORMED proteins, in the model plant Arabidopsis thaliana. In animals, much information is provided from the study of polarity in epithelial cells that exhibit basolateral and luminal apical polarities, separated by tight junctions. In this review, we summarize the similarities and differences of the polarization mechanisms between plants and animals and survey the main genetic approaches that have been used to characterize new genes involved in polarity establishment in plants, including the frequently used forward and reverse genetics screens as well as a novel chemical genetics approach that is expected to overcome the limitation of classical genetics methods.

Patterning in Arabidopsis root development is coordinated via a localized auxin concentration maximum in the root tip, requiring the regulated expression of specific genes. However, little is known about how hormone and gene expression patterning is generated., Using a variety of experimental data, we develop a spatiotemporal hormonal crosstalk model that describes the integrated action of auxin, ethylene and cytokinin signalling, the POLARIS protein, and the functions of PIN and AUX1 auxin transporters. We also conduct novel experiments to confirm our modelling predictions., The model reproduces auxin patterning and trends in wild-type and mutants; reveals that coordinated PIN and AUX1 activities are required to generate correct auxin patterning; correctly predicts shoot to root auxin flux, auxin patterning in the aux1 mutant, the amounts of cytokinin, ethylene and PIN protein, and PIN protein patterning in wild-type and mutant roots. Modelling analysis further reveals how PIN protein patterning is related to the POLARIS protein through ethylene signalling. Modelling prediction of the patterning of POLARIS expression is confirmed experimentally., Our combined modelling and experimental analysis reveals that a hormonal crosstalk network regulates the emergence of patterns and levels of hormones and gene expression in wild-type and mutants. [ABSTRACT FROM AUTHOR]

The plant hormone auxin is a key regulator of plant growth and development. Auxin levels are sensed and interpreted by distinct receptor systems that activate a broad range of cellular responses. The Auxin-Binding Protein1 (ABP1) that has been identified based on its ability to bind auxin with high affinity is a prime candidate for the extracellular receptor responsible for mediating a range of auxin effects, in particular, the fast non-transcriptional ones. Contradictory genetic studies suggested prominent or no importance of ABP1 in many developmental processes. However, how crucial the role of auxin binding to ABP1 is for its functions has not been addressed. Here, we show that the auxinbinding pocket of ABP1 is essential for its gain-of-function cellular and developmental roles. In total, 16 different abp1 mutants were prepared that possessed substitutions in the metal core or in the hydrophobic amino acids of the auxin-binding pocket as well as neutral mutations. Their analysis revealed that an intact auxin-binding pocket is a prerequisite for ABP1 to activate downstream components of the ABP1 signalling pathway, such as Rho of Plants (ROPs) and to mediate the clathrin association with membranes for endocytosis regulation. In planta analyses demonstrated the importance of the auxin binding pocket for all known ABP1-mediated postembryonic developmental processes, including morphology of leaf epidermal cells, root growth and root meristem activity, and vascular tissue differentiation. Taken together, these findings suggest that auxin binding to ABP1 is central to its function, supporting the role of ABP1 as auxin receptor. [ABSTRACT FROM AUTHOR]

Many aspects of development in the model plant Arabidopsis thaliana involve regulated distribution of the hormone auxin by the PIN-FORMED (PIN) family of auxin efflux carriers. The role of PIN-mediated auxin transport in other plants is not well understood, but studies in a wider range of species have begun to illuminate developmental mechanisms across land plants. In this review, I discuss recent progress in understanding the evolution of PIN-mediated auxin transport, and its role in development across the green plant lineage. I also discuss the idea that changes in auxin biology led to morphological novelty in plant development: currently available evidence suggests major innovations in auxin transport are rare and not associated with the evolution of new developmental mechanisms. [ABSTRACT FROM AUTHOR]

Timing of leaf abscission is an important trait for biomass production and seasonal acclimation in deciduous trees. The signaling leading to organ separation, from the external cue (decreasing photoperiod) to ethylene-regulated hydrolysis of the middle lamellae in the abscission zone, is only poorly understood. Data from annual species indicate that the formation of an auxin gradient spanning the abscission zone regulates the timing of abscission. We established an experimental system in Populus to induce leaf shedding synchronously under controlled greenhouse conditions in order to test the function of auxin in leaf abscission. Here, we show that exogenous auxin delayed abscission of dark-induced leaves over short and long distances and that a new auxin response maximum preceded the formation of an abscission zone. Several auxin transporters were down-regulated during abscission and inhibition of polar auxin transport delayed leaf shedding. Ethylene signaling was not involved in the regulation of these auxin transporters and in the formation of an abscission zone, but was required for the expression of hydrolytic enzymes associated with cell separation. Since exogenous auxin delayed abscission in absence of ethylene signaling auxin likely acts independently of ethylene signaling on cell separation. [ABSTRACT FROM AUTHOR]

In Agave tequilana, reproductive failure or inadequate flower development stimulates the formation of vegetative bulbils at the bracteoles, ensuring survival in a hostile environment. Little is known about the signals that trigger this probably unique phenomenon in agave species. Here we report that auxin plays a central role in bulbil development and show that the localization of PIN1-related proteins is consistent with altered auxin transport during this process. Analysis of agave transcriptome data led to the identification of the A. tequilana orthologue of PIN1 (denoted AtqPIN1) and a second closely related gene from a distinct clade reported as 'Sister of PIN1' (denoted AtqSoPIN1). Quantitative real-time reverse transcription--PCR (RT-qPCR) analysis showed different patterns of expression for each gene during bulbil formation, and heterologous expression of the A. tequilana PIN1 and SoPIN1 genes in Arabidopsis thaliana con- firmed functional differences between these genes. Although no free auxin was detected in induced pedicel samples, changes in the levels of auxin precursors were observed. Taken as a whole, the data support the model that AtqPIN1 and AtqSoPIN1 have co-ordinated but distinct functions in relation to auxin transport during the initial stages of bulbil formation. [ABSTRACT FROM AUTHOR]