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Photoperiod and temperature are vital environmental factors that regulate plant developmental processes. However, the roles of these factors in garlic bulb enlargement are unclear. In this report, responses of garlic bulb morphology and physiology to combinations of photoperiod (light/dark: 10/14 h, 12/12 h, 14/10 h) and temperature (light/dark: 25/18 °C, 30/20 °C) were investigated. For garlic cultivar G103, bulb characteristics, phytohormones (IAA, ABA, ZT, tZR, JA), allicin and phenolic acids (p-coumaric and p-hydroxybenzoic) were highest under a photoperiod of 14 h at 30 °C. Maximum GA was observed under 14 h + 30 °C for cv. G2011-5. Maximum caffeic, ferulic and vanillic acids were detected for cv. G2011-5 at 14 h + 30 °C, 12 h + 25 °C and 14 h + 25 °C, respectively. Flavonoids (myricetin, quercetin, kaempferol and apigenin) were not detected in this trial. This is the first report describing the impact of long periods of light duration and higher temperatures on garlic morphology, phytohormones, phenolic acids and allicin content.

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BACKGROUND: Geophytes possess specialized storage organs - bulbs, tubers, corms or rhizomes, which allow their survival during unfovarable periods and provide energy support for sprouting and sexual and vegetative reproduction. Bulbing and flowering of the geophyte depend on the combined effects of the internal and external factors, especially temperature and photoperiod. Many geophytes are extensively used in agriculture, but mechanisms of regulation of their flowering and bulbing are still unclear. RESULTS: Comparative morpho-physiological and transcriptome analyses and quantitative validation of gene expression shed light on the molecular regulation of the responses to vernalization in garlic, a typical bulbous plant. Long dark cold exposure of bulbs is a major cue for flowering and bulbing, and its interactions with the genetic makeup of the individual plant dictate the phenotypic expression during growth stage. Photoperiod signal is not involved in the initial nuclear and metabolic processes, but might play role in the later stages of development, flower stem elongation and bulbing. Vernalization for 12 weeks at 4 °C and planting in November resulted in flower initiation under short photoperiod in December-January, and early blooming and bulbing. In contrast, non-vernalized plants did not undergo meristem transition. Comparisons between vernalized and non-vernalized bulbs revealed ~ 14,000 differentially expressed genes. CONCLUSIONS: Low temperatures stimulate a large cascades of molecular mechanisms in garlic, and a variety of flowering pathways operate together for the benefit of meristem transition, annual life cycle and viable reproduction results.The circadian clock appears to play a central role in the transition of the meristem from vegetative to reproductive stage in bulbous plant, serving as integrator of the low-temperature signals and the expression of the genes associated with vernalization, photoperiod and meristem transition. The reserved photoperiodic pathway is integrated at an upstream point, possibly by the same receptors. Therefore, in bulb, low temperatures stimulate cascades of developmental mechanisms, and several genetic flowering pathways intermix to achieve successful sexual and vegetative reproduction.

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The photoperiod marks a varied set of behaviors in plants, including bulbing. Bulbing is controlled by inner signals, which can be stimulated or subdued by the ecological environment. It had been broadly stated that phytohormones control the plant development, and they are considered to play a significant part in the bulb formation. The past decade has witnessed significant progress in understanding and advancement about the photoperiodic initiation of bulbing in plants. A noticeable query is to what degree the mechanisms discovered in bulb crops are also shared by other species and what other qualities are also dependent on photoperiod. The FLOWERING LOCUS T (FT) protein has a role in flowering; however, the FT genes were afterward reported to play further functions in other biological developments (e.g., bulbing). This is predominantly applicable in photoperiodic regulation, where the FT genes seem to have experienced significant development at the practical level and play a novel part in the switch of bulb formation in Alliums. The neofunctionalization of FT homologs in the photoperiodic environments detects these proteins as a new class of primary signaling mechanisms that control the growth and organogenesis in these agronomic-related species. In the present review, we report the underlying mechanisms regulating the photoperiodic-mediated bulb enlargement in Allium species. Therefore, the present review aims to systematically review the published literature on the bulbing mechanism of Allium crops in response to photoperiod. We also provide evidence showing that the bulbing transitions are controlled by phytohormones signaling and FT-like paralogues that respond to independent environmental cues (photoperiod), and we also show that an autorelay mechanism involving FT modulates the expression of the bulbing-control gene. Although a large number of studies have been conducted, several limitations and research gaps have been identified that need to be addressed in future studies.

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TERMINAL FLOWER1 (TFL1) is a key gene for maintenance of vegetative and inflorescence indeterminacy and architecture. In onion, flowering and bulbing are two distinct developmental phases, each under complex environmental regulatory control. We have identified two CEN/TFL1-like genes from onion designated as AcTFL1 and AcCEN1. AcTFL1 is expressed during bulbing and inflorescence development with expression increasing with indeterminate growth of the umbel and the bulb suggesting possible conservation of function. Increase in AcTFL1 expression during umbel growth is associated with a simultaneous reduction in expression of AcLFY. Expression of AcTFL1 within the bulb is lowest in the outermost layers and highest in the innermost (youngest) layers. Bulb storage at room temperature or in cold leads to a gradual reduction in AcTFL1 levels in the meristem-containing tissues, the decrease being faster in the variety not requiring vernalization. Constitutive expression of AcTFL1, but not AcCEN1 complements the Arabidopsis tfl1-14 mutant and delays flowering in wild type suggesting conservation of the AcTFL1 function even in the distantly related Arabidopsis. Taken together, AcTFL1 appears to be the functional counterpart of TFL1 and regulates indeterminate growth of the umbel inflorescence as well as bulb development in onion.

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This research was conducted to evaluate the effect of temperature and photoperiod treatments on the bolting and bulb formation of three local garlic cultivars (cvs) in two consecutive years. Naturally vernalized plants of cvs G107, G025 and G064 were transplanted into growth chambers and subjected to various combinations of temperature [T15/10, 15°C/10°C; T20/15, 20°C/15°C and T25/18, 25°C/18°C (day/night)] and photoperiod (L8, 8 h and L14,14 h) treatments. Plant growth, endogenous phytohormone and methyl jasmonate (MeJA) levels, along with the bolting and yield of garlic were evaluated. The experimental results from two consecutive years indicated that higher temperature (20°C or 25°C) and longer photoperiod (14 h) treatments significantly enhanced the garlic bolting, bulbing and cloving with a shorter growth period and a higher bulb weight. Moreover, the endogenous phytohormone and MeJA levels in the test plants were significantly increased by the higher temperature (25°C for the phytohormone level; 20°C for the MeJA level) and longer photoperiod [14 h, except for abscisic acid (ABA), which had the highest level at 8 h] conditions and were decreased by the lowest test temperature (15°C) and shorter photoperiod (8 h, except for ABA) conditions. This response coincided with that of the bulbing index, bolting rate, growth period and bulb weight. In addition, plants treated under the conditions of 20°C/15°C-14 h and 25°C/18°C-14 h produced the highest phytohormone levels (except for ABA) for cvs G025 and G064, respectively, and showed the best bolting and bulbing behavior. It is reasonable to assume that endogenous phytohormone (especially gibberellic acid) and MeJA levels are highly related to garlic bolting and bulbing, which might lead to the different responses of the three studied cultivars to the combination of temperature and photoperiod treatments. Furthermore, cvs G107 and G025 bolt well and have better bulb formation under 20°C-14 h conditions, while the conditions of 25°C-14 h are critical for the bolting and bulbing of cv. G064.

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A protocol for obtaining bulbs via in vitro organogenesis was developed for tarda tulip (Tulipa tarda Stapf). Scale explants were obtained from bulbs formed at the base of seedlings or from adventitious bulbs that developed from callus tissue forming on stolons or on germinating seeds. Some explants were subjected to chilling at 5°C for 12 wk. The culture media contained 3 or 6% sucrose and was supplemented with either no growth regulators, either 0.5 µM 6-benzyl-aminopurine (BAP) or 18.9 or 94.6 µM abscisic acid (ABA). Cultures were maintained in the dark at 20°C. Callus tissue developed mainly on media without growth regulators or with BAP. Callus was formed from up to 96% of explants derived from non-chilled adventitious bulbs that were treated with 3% sucrose and 0.5 µM BAP. Less callus was formed from chilled explants compared with non-chilled explants. Newly formed adventitious bulbs appeared on the explants via direct and indirect organogenesis. The media with BAP promoted the formation of adventitious bulbs at a rate of 56-92% from non-chilled explants, whereas a maximum rate of 36% was observed from chilled explants. ABA inhibited the induction of adventitious bulbs and callus. The adventitious bulbs obtained in these experiments contained a meristem, which was evidence that they had developed properly.

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Protocolos eficientes de crescimento de ápices caulinares de alho (Allium sativum L.) e posterior bulbificação in vitro são importantes para limpeza clonal e manutenção da fidelidade genética. O trabalho teve como objetivo avaliar os efeitos de tipos e concentrações de reguladores de crescimento sobre a morfogênese de plantas de alho in vitro. Ápices caulinares com até dois primórdios foram excisados de bulbilhos de alho da cv. 'Jonas' e submetidos ao cultivo in vitro em meio de cultura suplementado de ácido indolacético (0; 1,07; 2,69 e 5,37µM), ácido indolbutírico (0; 0,49; 0,98 e 2,46µM), ácido naftalenoacético (0; 1,07; 2,69 e 5,37µM), ácido jasmônico (0; 1,0; 5,0 e 10,0µM) e ácido abscísico (0; 0,38; 1,89; e 3,78µM). A concentração de 1,07µM de ácido naftalenoacético aplicado ao meio de cultura promoveu incrementos na maioria das variáveis analisadas. O ácido jasmônico induziu a formação de bulbos de alho in vitro, embora tenha apresentado performance inferior ao verificado com o uso de ANA. Por outro lado, a adição de ácido abscísico no meio de cultura inibiu o crescimento de plantas, porém, não impediu a formação de bulbos, sobretudo em concentrações reduzidas. De um modo geral, as variáveis número de bulbos e porcentagem de bulbificação diminuiram com o uso de concentrações elevadas dos reguladores de crescimento testados. Entre os reguladores de crescimento de plantas, o ANA apresenta maior efeito na morfogênese in vitro de plantas de alho, entretanto, o ácido jasmônico e o ABA também apresentam potencial para induzir a formação de bulbos de alho in vitro como o ANA.

Efficient protocols for garlic (Allium sativum L.) shoot tips growth and later in vitro bulbing are significant for pathogen removal and maintenance of genetic fidelity. The objective of this study was to assess the effects of different types and concentrations of growth regulators on in vitro morphogenesis of garlic plants. Shoot meristems with up to two primordial were excised from garlic bulbils (cv. 'Jonas') and cultivated in vitro in culture media supplemented with indoleacetic acid (0; 1,07; 2,69 e 5,37µM), indolbutyric acid (0; 0,49; 0,98 e 2,46µM), naphthaleneacetic acid (0; 1,07; 2,69 e 5,37µM), and abscisic acid (0; 0,38; 1,89; e 3,78µM). The concentration of 1,07µM indoleacetic acid applied to the culture medium promoted increases in most variables analyzed. Jasmonic acid induced formation of garlic bulbs in vitro, although it showed lower performance than that verified with the use of NAA. On the other hand, addition of abscisic acid in culture media inhibited plant growth. However, it did not impede the formation of bulbs, especially when in reduced concentrations. Generally speaking, variables such as number of bulbs and rate of bulbing decreased with the use of high concentrations of the assessed growth regulators. Among plant growth regulators, NAA showed a stronger effect on in vitro morphogenesis of garlic plants. Nonetheless, jasmonic acid and ABA also showed potential to induce formation of garlic bulbs in vitro such as NAA.

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The tissue-specific and developmental expression of histone 2A was studied in onion (Allium cepa'Robusta'), using northern blots. Histone 2A expression was enriched in basal tissues, particularly in the inner, meristematically active parts of bulbs. The expression was assessed during a time course of bulb development, dormancy onset and post-harvest sprouting in field-grown material. During bulb development histone 2A expression in the inner bulb declined rapidly during bulb ripening, reaching a minimum with the onset of dormancy. During post-harvest storage, expression increased slowly, reaching a peak in the spring, coinciding with the first observed sprout emergence. It was concluded that in onion, as in other plant systems, histone 2A expression is linked to cell division and dormancy level, the peak in expression during post-harvest storage indicating the time of dormancy breakage. In cultivars where post-harvest sprouting occurred much earlier or much later than in 'Robusta', this expression peak occurred at about the same time of year, regardless of sprouting time. It was concluded that differences in storage longevity between cultivars were not due to differing times of dormancy breakage. Factors controlling the rate of sprout emergence post-dormancy are likely to be major determinants of storage capacity.

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Twenty-three patients with pulstile bulging on the subclavian (14) and carotid triangles of the neck (9) were investigated with duplex ultrasonography and CT angiography to determine their causes. The causes of pulsatile bulging were innominate tortuosity (14) at right subclavian triangle and prominant carotid sinus (4), common carotid artery tortuosity (2), common carotid artery pseudoaneurysm (1), and carotid artery thrombosis (1) at carotid triangle.

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