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• Clonal growth of plants is attained by a number of morphologically different organs (e.g. stolons, rhizomes, and roots), which are not functionally equivalent. Consequently, these clonal growth organ (CGO) types can determine functional traits that are associated with clonality, although little is known about their evolutionary flexibility or the constraining role they play on clonal traits. • We investigated the rates of evolutionary change by which individual CGOs are acquired and lost using a set of 2652 species of Central European flora. Furthermore, we asked how these individual CGOs constrain functionally relevant clonal traits, such as lateral spread, number of offspring, and persistence of connections. • We show that plants can easily switch in evolution among individual types of CGO and between clonal and nonclonal habits. However, not all these transitions are equally probable. Namely, stem-based clonal growth and root-based clonal growth constitute evolutionarily separate forms of clonal growth. • Clonal traits are strongly constrained by individual CGO types. Specifically, fast lateral spread is attained by stolons or hypogeogenous rhizomes, and persistent connections are attained by all rhizome types. However, the ease with which clonal organs appear and disappear in evolution implies that plants can overcome these constraints by adjusting their morphologies.

The enrichment with Trifolium repens L. (white clover), could fulfill an ecological and productive function in humid-temperate silvopastoral systems in South America. The objective was to describe the morphogenetic, structural and functional changes of two white clover large-leafed cultivars under simultaneous shading and phosphorus shortage during establishment. A pot trial designed in 3 randomized complete blocks was carried out in sub-subdivided plots. The main plot was the shading treatment (4 levels: full sun = 0% and 30%, 60% and 90% of shading (shade), sub-plots were 2 cultivars (Junín and El Lucero) and sub-sub-plots, were 2 phosphorous conditions (without P− and with added P+). At final harvest the number of primary and secondary stolons and the length of their internodes, the number of leaves and length of petioles were counted. The light threshold was on 60% of shading; mean total daily PAR > 15.3 mol m −2  day −1 . This light threshold was the same under P+ or P−. The number of stolons and length of internodes remained between full sun and 60% shade but the length of petioles increased. The lower number of expanding leaves was compensated by a lower number of senescent ones that mantained the living leaves and leaf area of the plants up to 60% of shading. P shortage did not limit the number of primary or secondary stolons, the number of leaves or petiole length, so the horizontal or vertical occupation was slightly affected. Both cultivars responded similarly. White clover would be a promising species for these systems.

1. This account presents comprehensive information on the biology of Phragmites australis (Cav.) Trin. ex Steud. (P. communis Trin.; common reed) that is relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles: distribution, habitat, communities, responses to biotic factors and to the abiotic environment, plant structure and physiology, phenology, floral and seed characters, herbivores and diseases, as well as history including invasive spread in other regions, and conservation. 2. Phragmites australis is a cosmopolitan species native to the British flora and widespread in lowland habitats throughout, from the Shetland archipelago to southern England. It is widespread throughout Ireland and is native in the Channel Islands. Native populations occur naturally in temperate zones and on every continent except Antarctica. Some populations in Australia and North America have been introduced from elsewhere and have become naturalized, and in North America, some of these are known to be invasive where they compete with native local populations of P. australis. Typical habitats in Britain range from shallow still water along waterbody edges to marshlands, saltmarshes and drier habitat on slopes up to 470 m above sea level. Additional habitats outside Britain are springs in arid areas, riverine lowlands (— 5 m above sea level) and groundwater seepage points up to 3600 m above sea level. Although it occurs on a wide range of substrates and can tolerate pH from 2.5 to 9.8, in Britain it prefers pH >4.5 and elsewhere it thrives in mildly acidic to mildly basic conditions (pH 5.5-7.5). The species plays a pivotal role in the successional transition from open water to woodland. 3. Phragmites australis is a tall, helophytic, wind-pollinated grass with annual shoots up to 5 m above-ground level from an extensive system of rhizomes and stolons. A single silky inflorescence develops at the end of each fertile stem and produces 500—2000 seeds. The plant is highly variable genetically and morphologically. 4. Expansion of established populations is mainly through clonal growth of the horizontal rhizome system and ground-surface stolons, while new populations can establish from rhizomes, stem fragments and seeds. Shoots generally emerge in spring, with timing determined primarily by physiology that is mediated by external conditions (e.g. local climate including frost). 5. Many populations in the British Isles have experienced some decline over the past two decades and there is concern that there might be further losses along the east coast as sea level rises. There have recently also been localized expansions, especially in highly modified habitats, where P. australis reedbeds have been planted as wildlife habitat, rehabilitated mineral and gravel beds, and bioremediation filter beds for industrial and transport infrastructure. Native populations outside Britain also demonstrate both types of trend: they are declining in many parts of Western Europe and North America, yet also colonize many disturbed, ruderal habitats (e.g. the edges of agricultural fields and motorways) throughout its native and non-native range and can form 'weedy' monodominant populations (e.g. in Australia and China).

• Plant architecture is central in determining crop yield. In the short-day species strawberry, a crop vegetatively propagated by daughter-plants produced by stolons, fruit yield is further dependent on the trade-off between sexual reproduction (fruits) and asexual reproduction (daughter-plants). Both are largely dependent on meristem identity, which establishes the development of branches, stolons and inflorescences. • Floral initiation and plant architecture are modulated by the balance between two related proteins, FLOWERING LOCUS T (FT) and TERMINAL FLOWER 1 (TFL1). We explored in woodland strawberry the role of the uncharacterised FveFT2 and FveFT3 genes and of the floral repressor FveTFL1 through gene expression analyses, grafting and genetic transformation (overexpression and gene editing). • We demonstrate the unusual properties of these genes. FveFT2 is a nonphotoperiodic florigen permitting short-day (SD) flowering and FveTFL1 is the long-hypothesised long-day systemic antiflorigen that contributes, together with FveFT2, to the photoperiodic regulation of flowering. We additionally show that FveFT3 is not a florigen but promotes plant branching when overexpressed, that is likely to be through changing axillary meristem fate, therefore resulting in a 3.5-fold increase in fruit yield at the expense of stolons. • We show that our findings can be translated into improvement of cultivated strawberry in which FveFT2 overexpression significantly accelerates flowering.

MicroRNA156 (miR156) functions in maintaining the juvenile phase in plants. However, the mobility of this microRNA has not been demonstrated. So far, only three microRNAs, miR399, miR395, and miR172, have been shown to be mobile. We demonstrate here that miR156 is a potential graft-transmissible signal that affects plant architecture and tuberization in potato (Solanum tuberosum). Under tuber-noninductive (long-day) conditions, miR156 shows higher abundance in leaves and stems, whereas an increase in abundance of miR156 has been observed in stolons under tuber-inductive (short-day) conditions, indicative of a photoperiodic control. Detection of miR156 in phloem cells of wild-type plants and mobility assays in heterografts suggest that miR156 is a graft-transmissible signal. This movement was correlated with changes in leaf morphology and longer trichomes in leaves. Overexpression of miR156 in potato caused a drastic phenotype resulting in altered plant architecture and reduced tuber yield. miR156 overexpression plants also exhibited altered levels of cytokinin and strigolactone along with increased levels of LONELY GUY1 and StCyclin D3.1 transcripts as compared with wild-type plants. RNA ligase-mediated rapid amplification of complementary DNA ends analysis validated SQUAMOSA PROMOTER BINDING-LIKE3 (StSPL3), StSPL6, StSPL9, StSPL13, and StLIGULELESS1 as targets of miR156. Gel-shift assays indicate the regulation of miR172 by miR156 through StSPL9. miR156-resistant SPL9 overexpression lines exhibited increased miR172 levels under a short-day photoperiod, supporting miR172 regulation via the miR156-SPL9 module. Overall, our results strongly suggest that miR156 is a phloem-mobile signal regulating potato development.

As a perennial warm-season turfgrass species with great economic value, bermudagrass (Cynodon dactylon L.) simultaneously has three types of stems: shoot, stolon, and rhizome. However, molecular mechanisms underlying the specialization of the three types of stems remain poorly understood. In this study, the metabolome differences among the three types of stems were analyzed and compared through untargeted metabolomic profiling in combination with transcriptome-wide analyses of the genes participating in the metabolic pathways. A total of 949 metabolites were identified in the three stems, whereas 303, 473, and 330 metabolites were differentially accumulated between shoots and stolons, shoots and rhizomes, and stolons and rhizomes, respectively. Sugars and phenylpropanoids were two enriched categories of metabolites showing preferential accumulation in the three types of stems. Transcriptome and RT-qPCR analyses indicated that gene expression of key enzymes catalyzing the synthesis and transformation of sugars and phenylpropanoids, especially glucose-1-phosphate adenylyltransferase, starch synthase, and phenylalanine ammonia-lyase, were delicately regulated to maintain the sugar-starch and lignin-flavonoid homeostasis in the three stems. The results of this study not only expanded our understanding of metabolism regulation in bermudagrass, but also laid a foundation for molecular mechanism study of stem specialization in this glamorous plant species.

Core Ideas WinRHIZO technology may provide a method to analyze stolon morphology of grasses. The scanned length closely predicted the manually measured one. WinRHIZO technology overestimated diameter when the whole stolon was scanned. WinRHIZO, a root‐measuring system, may provide an alternative, reliable, and fast method to analyze stolon morphology of grass species. This study evaluated the possibility to use WinRHIZO technology to measure total length and average diameter of bermudagrass [Cynodon dactylon (L.) Pers.] stolons. The length and diameter of 70 stolons collected from four turf‐type cultivars and a wild bermudagrass were measured with a ruler (length) and caliper (diameter), and using WinRHIZO technology, a scanner‐based image analysis system. The scanned length closely predicted the manually measured one and can be successfully used to determine stolon length in samples with a significant amount of biomass. WinRHIZO technology overestimated diameter values when the whole stolon was scanned, while the diameter prediction was more precise when nodes were removed and only internodes were measured.

To uncover the nature of various kinds of stolons of Utricularia subgenus Polypompholyx (Lentibulariaceae) we studied branching of stolons by scanning electron microscopy, statistically investigated correlations of stolon types and other traits across 56 species, and evaluated seedling development and process morphological aspects. Some results were compared to the sister genera Pinguicula and Genlisea . A key to nine stolon types in Polypompholyx is provided. Predominant stolon types were rhizoids, runner stolons with rhizoids on nodes, and runner stolons without rhizoids on nodes but with bladders on internodes. Stolon types were taxonomically relevant and correlated to the distribution/climate. They obviously diverged with speciation events in Australia. Examined seedlings of Genlisea and Polypompholyx showed similar developmental patterns. Stolons were homologous to traps and leaves. Selected subterranean organs contained specific but similar process combinations of roots, shoots and/or leaves. We assume the Genlisea - Utricularia ancestor trap included processes of a Pinguicula root and leaf.

Potato (Solanum tuberosum L.) can reproduce sexually through flowering and asexually through tuberization. While tuberization has been thoroughly studied, little research has been done on potato flowering. Flower bud development in the strictly short-day tuberizing S. tuberosum group Andigena is impaired under short-day conditions. This impaired development may indicate that tuberization negatively influences flowering. Here, we determine how tuberization affects flower bud development. To find out whether the absence of tubers improves flowering, we prevented tuberization by: (i) grafting potato scions onto wild potato rootstocks, which were unable to form tubers; (ii) removing stolons, the underground structures on which tubers form; and (iii) using plants that were silenced in the tuberization signal StSP6A. Additionally, transgenic plants with increased StSP6A expression were used to determine if flower bud development was impaired. The absence of a tuber sink alone did not accelerate flower bud development, nor did it allow more plants to reach anthesis (open flowering stage) or have more open flowers. Interestingly, reducing StSP6A expression improved flower bud development, and increasing expression impaired it. Our results show that flower bud development in potato is repressed by the tuberization signal StSP6A, and not by competition with the underground tuber sink.

Background Bermudagrass ( Cynodon dactylon L.) is an important warm-season turfgrass species with well-developed stolons, which lay the foundation for the fast propagation of bermudagrass plants through asexual clonal growth. However, the growth and development of bermudagrass stolons are still poorly understood at the molecular level. Results In this study, we comprehensively analyzed the acetylation and succinylation modifications of proteins in fast-growing stolons of the bermudagrass cultivar Yangjiang. A total of 4657 lysine acetylation sites on 1914 proteins and 226 lysine succinylation sites on 128 proteins were successfully identified using liquid chromatography coupled to tandem mass spectrometry, respectively. Furthermore, 78 proteins and 81 lysine sites were found to be both acetylated and succinylated. Functional enrichment analysis revealed that acetylated proteins regulate diverse reactions of carbohydrate metabolism and protein turnover, whereas succinylated proteins mainly regulate the citrate cycle. These results partly explained the different growth disturbances of bermudagrass stolons under treatment with sodium butyrate and sodium malonate, which interfere with protein acetylation and succinylation, respectively. Moreover, 140 acetylated proteins and 42 succinylated proteins were further characterized having similarly modified orthologs in other grass species. Site-specific mutations combined with enzymatic activity assays indicated that the conserved acetylation of catalase and succinylation of malate dehydrogenase both inhibited their activities, further implying important regulatory roles of the two modifications. Conclusion In summary, our study implied that lysine acetylation and succinylation of proteins possibly play important regulatory roles in the fast growth of bermudagrass stolons. The results not only provide new insights into clonal growth of bermudagrass but also offer a rich resource for functional analyses of protein lysine acetylation and succinylation in plants.