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Floral transition under drought conditions is accelerated by enabling ABA-dependent up-regulation of the florigen genes. Modulation of the transition to flowering plays an important role in the adaptation to drought. The drought-escape (DE) response allows plants to adaptively shorten their life cycle to make seeds before severe stress leads to death. However, the molecular basis of the DE response is unknown. A screen of different Arabidopsis (Arabidopsis thaliana) flowering time mutants under DE-triggering conditions revealed the central role of the flower-promoting gene GIGANTEA (GI) and the florigen genes FLOWERING LOCUS T (FT) and TWIN SISTER OF FT (TSF) in the DE response. Further screens showed that the phytohormone abscisic acid is required for the DE response, positively regulating flowering under long-day conditions. Drought stress promotes the transcriptional up-regulation of the florigens in an abscisic acid- and photoperiod-dependent manner, so that early flowering only occurs under long days. Along with the florigens, the floral integrator SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1 is also up-regulated in a similar fashion and contributes to the activation of TSF. The DE response was recovered under short days in the absence of the floral repressor SHORT VEGETATIVE PHASE or in GI-overexpressing plants. Our data reveal a key role for GI in connecting photoperiodic cues and environmental stress independently from the central FT/TSF activator CONSTANS. This mechanism explains how environmental cues may act upon the florigen genes in a photoperiodically controlled manner, thus enabling plastic flowering responses.

We studied the mechanical process of seed pods opening in Bauhinia variegate and found a chirality-creating mechanism, which turns an initially flat pod valve into a helix. We studied configurations of strips cut from pod valve tissue and from composite elastic materials that mimic its structure. The experiments reveal various helical configurations with sharp morphological transitions between them. Using the mathematical framework of \"incompatible elasticity,\" we modeled the pod as a thin strip with a flat intrinsic metric and a saddle-like intrinsic curvature. Our theoretical analysis quantitatively predicts all observed configurations, thus linking the pod's microscopic structure and macroscopic conformation. We suggest that this type of incompatible strip is likely to play a role in the self-assembly of chiral macromolecules and could be used for the engineering of synthetic self-shaping devices.

Double fertilization is the defining characteristic of flowering plants. However, the molecular mechanisms regulating the fusion of one sperm with the egg and the second sperm with the central cell are largely unknown. We show that gamete interactions in Arabidopsis depend on small cysteine-rich EC1 (EGG CELL 1) proteins accumulating in storage vesicles of the egg cell. Upon sperm arrival, EC1-containing vesicles are exocytosed. The sperm endomembrane system responds to exogenously applied EC1 peptides by redistributing the potential gamete fusogen HAP2/GCS1 (HAPLESS 2/GENERATIVE CELL SPECIFIC 1) to the cell surface. Furthermore, fertilization studies with ec1 quintuple mutants show that successful male-female gamete interactions are necessary to prevent multiple-sperm cell delivery. Our findings provide evidence that mutual gamete activation, regulated exocytosis, and sperm plasma membrane modifications govern flowering plant gamete interactions.

The pollen tube is a cellular protuberance formed by the pollen grain, or male gametophyte, in flowering plants. Its principal metabolic activity is the synthesis and assembly of cell wall material, which must be precisely coordinated to sustain the characteristic rapid growth rate and to ensure geometrically correct and efficient cellular morphogenesis. Unlike other model species, the cell wall of the Arabidopsis (Arabidopsis thaliana) pollen tube has not been described in detail. We used immunohistochemistry and quantitative image analysis to provide a detailed profile of the spatial distribution of the major cell wall polymers composing the Arabidopsis pollen tube cell wall. Comparison with predictions made by a mechanical model for pollen tube growth revealed the importance of pectin deesterification in determining the cell diameter. Scanning electron microscopy demonstrated that cellulose microfibrils are oriented in near longitudinal orientation in the Arabidopsis pollen tube cell wall, consistent with a linear arrangement of cellulose synthase CESA6 in the plasma membrane. The cellulose label was also found inside cytoplasmic vesicles and might originate from an early activation of cellulose synthases prior to their insertion into the plasma membrane or from recycling of short cellulose polymers by endocytosis. A series of strategic enzymatic treatments also suggests that pectins, cellulose, and callose are highly cross linked to each other.

Since the discovery of miRNAs in plants it has become clear that they are central to the regulation of many aspects of plant development and responses to the environment. miR172 regulates expression of a small group of AP2-like transcription factors in an evolutionarily ancient interaction. miR172 functions in regulating the transitions between developmental stages and in specifying floral organ identity. These two roles are conserved across monocotyledons and dicotyledons. Investigations into the roles of miR172 and its targets in phase changes in the model plant Arabidopsis have illustrated that this process is governed by complex regulatory systems. In addition to its conserved roles, miR172 has also acquired specialized species-specific functions in other aspects of plant development such as cleistogamy and tuberization.

Precise timing of CONSTANS (CO) gene expression is necessary for day-length discrimination for photoperiodic flowering. The FLAVIN-BINDING, KELCH REPEAT, F-BOX 1 (FKF1), and GIGANTEA (GI) proteins regulate CO transcription in ARABIDOPSIS: We demonstrate that FKF1 and GI proteins form a complex in a blue-light-dependent manner. The timing of this interaction regulates the timing of daytime CO expression. FKF1 function is dependent on GI, which interacts with a CO repressor, CYCLING DOF FACTOR 1 (CDF1), and controls CDF1 stability. GI, FKF1, and CDF1 proteins associate with CO chromatin. Thus, the FKF1-GI complex forms on the CO promoter in late afternoon to regulate CO expression, providing a mechanistic view of how the coincidence of light with circadian timing regulates photoperiodic flowering.

In plants, seasonal changes in day length are perceived in leaves, which initiate long-distance signaling that induces flowering at the shoot apex. The identity of the long-distance signal has yet to be determined. In Arabidopsis, activation of FLOWERING LOCUS T (FT) transcription in leaf vascular tissue (phloem) induces flowering. We found that FT messenger RNA is required only transiently in the leaf. In addition, FT fusion proteins expressed specifically in phloem cells move to the apex and move long distances between grafted plants. Finally, we provide evidence that FT does not activate an intermediate messenger in leaves. We conclude that FT protein acts as a long-distance signal that induces Arabidopsis flowering.

In higher plants, timely degradation of tapetai cells, the innermost sporophytic cells of the anther wall layer, is a prerequisite for the development of viable pollen grains. However, relatively little is known about the mechanism underlying programmed tapetai cell development and degradation. Here, we report a key regulator in monocot rice (Oryza sauva), PERSISTANT TAPETAL CELLI (PTCl), which controls programmed tapetal development and functional pollen formation. The evolutionary significance of PTCl was revealed by partial genetic complementation of the homologous mutation MALE STERILITY1 (MSI) in the dicot Arabidopsis (Arabidopsis thaliana). PTCl encodes a PHD-finger (for plant homeodomain) protein, which is expressed specifically in tapetai cells and microspores during anther development in stages 8 and 9, when the wild-type tapetai cells initiate a typical apoptosis-like cell death. Even though ptcl mutants show phenotypic similarity to msl in a lack of tapetai DNA fragmentation, delayed tapetai degeneration, as well as abnormal pollen wall formation and aborted microspore development, the ptcl mutant displays a previously unreported phenotype of uncontrolled tapetai proliferation and subsequent commencement of necrosis-like tapetai death. Microarray analysis indicated that 2,417 tapetum-and microspore-expressed genes, which are principally associated with tapetai development, degeneration, and pollen wall formation, had changed expression in ptcl anthers. Moreover, the regulatory role of PTCl in anther development was revealed by comparison with MS2 and other rice anther developmental regulators. These findings suggest a diversified and conserved switch of PTCl /MSI in regulating programmed male reproductive development in both dicots and monocots, which provides new insights in plant anther development.

Self-incompatibility in flowering plants prevents inbreeding and promotes outcrossing to generate genetic diversity. In Solanaceae, a multiallelic gene, S-locus F-box (SLF), was previously shown to encode the pollen determinant in self-incompatibility. It was postulated that an SLF allelic product specifically detoxifies its non-self S-ribonucleases (S-RNases), allelic products of the pistil determinant, inside pollen tubes via the ubiquitin-26S-proteasome system, thereby allowing compatible pollinations. However, it remained puzzling how SLF, with much lower allelic sequence diversity than S-RNase, might have the capacity to recognize a large repertoire of non-self S-RNases. We used in vivo functional assays and protein interaction assays to show that in Petunia, at least three types of divergent SLF proteins function as the pollen determinant, each recognizing a subset of non-self S-RNases. Our findings reveal a collaborative non-self recognition system in plants.

Florigen, the mobile signal that moves from an induced leaf to the shoot apex and causes flowering, has eluded identification since it was first proposed 70 years ago. Understanding the nature of the mobile flowering signal would provide a key insight into the molecular mechanism of floral induction. Recent studies suggest that the Arabidopsis FLOWERING LOCUS T (FT) gene is a candidate for encoding florigen. We show that the protein encoded by Hd3a, a rice ortholog of FT, moves from the leaf to the shoot apical meristem and induces flowering in rice. These results suggest that the Hd3a protein may be the rice florigen.