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Following tuber induction, potato tubers undergo a period of dormancy during which visible bud growth is inhibited. The length of the dormancy period is under environmental, physiological and hormonal control. Sucrose availability is one prerequisite for bud break. In the absence of sucrose, no bud break occurs. Thus, sucrose is likely to serve as nutrient and signal molecule at the same time. The mode of sucrose sensing is only vaguely understood, but most likely involves trehalose-6-phosphate and SnRK1 signalling networks. This conclusion is supported by the observation that ectopically manipulation of trehalose-6-phosphate levels influences the length of the dormancy period. Once physiological competence is achieved, sprouting is controlled by the level of phytohormones. Two phytohormones, ABA and ethylene, are supposed to suppress tuber sprouting; however, the exact role of ethylene remains to be elucidated. Cytokinins and gibberellins are required for bud break and sprout growth, respectively. The fifth classical phytohormone, auxin, seems to play a role in vascular development. During the dormancy period, buds are symplastically isolated, which changes during bud break. In parallel to the establishment of symplastic connectivity, vascular tissue develops below the growing bud most likely to support the outgrowing sprout with assimilates mobilised in parenchyma cells. Sprouting leads to major quality losses of stored potato tubers. Therefore, control of tuber sprouting is a major objective in potato breeding. Although comparative transcriptome analysis revealed a large number of genes differentially expressed in growing versus dormant buds, no master-regulator of potato tuber sprouting has been identified so far.

Trehalose-6-phosphate (T6P) is a signaling metabolite that regulates carbon metabolism, developmental processes, and growth in plants. In Arabidopsis (Arabidopsis thaliana), T6P signaling is, at least in part, mediated through inhibition of the SNF1-related protein kinase SnRKl. To investigate the role of T6P signaling in a heterotrophic, starch-accumulating storage organ, transgenic potato (Solatium tuberosum) plants with altered T6P levels specifically in their tubers were generated. Transgenic lines with elevated T6P levels (B33-TPS, expressing Escherichia coli osmoregulatory trehalose synthesis A [OtsA], which encodes a T6P synthase) displayed reduced starch content, decreased ATP contents, and increased respiration rate diagnostic for high metabolic activity. On the other hand, lines with significantly reduced T6P (B33-TPP, expressing E. coli OtsB, which encodes a T6P phosphatase) showed accumulation of soluble carbohydrates, hexose phosphates, and ATP, no change in starch when calculated on a fresh weight basis, and a strongly reduced tuber yield. [¹⁴C] Glucose feeding to transgenic tubers indicated that carbon partitioning between starch and soluble carbohydrates was not altered. Transcriptional profiling of B33-TPP tubers revealed that target genes of SnRKl were strongly up-regulated and that T6P inhibited potato tuber SnRKl activity in vitro. Among the SnRKl target genes in B33-TPP tubers, those involved in the promotion of cell proliferation and growth were down-regulated, while an inhibitor of cell cycle progression was up-regulated. T6P-accumulating tubers were strongly delayed in sprouting, while those with reduced T6P sprouted earlier than the wild type. Early sprouting of B33-TPP tubers correlated with a reduced abscisic acid content. Collectively, our data indicate that T6P plays an important role for potato tuber growth.

Key messageIn this study we show that expression of the Arabidopsis ABF4 gene in potato increases tuber yield under normal and abiotic stress conditions, improves storage capability and processing quality of the tubers, and enhances salt and drought tolerance.Potato is the third most important food crop in the world. Potato plants are susceptible to salinity and drought, which negatively affect crop yield, tuber quality and market value. The development of new varieties with higher yields and increased tolerance to adverse environmental conditions is a main objective in potato breeding. In addition, tubers suffer from undesirable sprouting during storage that leads to major quality losses; therefore, the control of tuber sprouting is of considerable economic importance. ABF (ABRE-binding factor) proteins are bZIP transcription factors that regulate abscisic acid signaling during abiotic stress. ABF proteins also play an important role in the tuberization induction. We developed transgenic potato plants constitutively expressing the Arabidopsis ABF4 gene (35S::ABF4). In this study, we evaluated the performance of 35S::ABF4 plants grown in soil, determining different parameters related to tuber yield, tuber quality (carbohydrates content and sprouting behavior) and tolerance to salt and drought stress. Besides enhancing salt stress and drought tolerance, constitutive expression of ABF4 increases tuber yield under normal and stress conditions, enhances storage capability and improves the processing quality of the tubers.

Various transcriptional networks and plant hormones have been implicated in controlling different aspects of potato tuber formation. Due to its broad impact on many plant developmental processes, a role for auxin in tuber initiation has been suggested but never fully resolved. Here, auxin concentrations were measured throughout the plant prior to and during the process of tuber formation. Auxin levels increase dramatically in the stolon prior to tuberization and remain relatively high during subsequent tuber growth, suggesting a promoting role for auxin in tuber formation. Furthermore, in vitro tuberization experiments showed higher levels of tuber formation from axillary buds of explants where the auxin source (stolon tip) had been removed. This phenotype could be rescued by application of auxin on the ablated stolon tips. In addition, a synthetic strigolactone analogue applied on the basal part of the stolon resulted in fewer tubers. The experiments indicate that a system for the production and directional transport of auxin exists in stolons and acts synergistically with strigolactones to control the outgrowth of the axillary stolon buds, similar to the control of above-ground shoot branching.

Dual florigen response in potatoes The seasonality of plant developmental processes such as flowering and tuber formation is dependent largely on changes in day length. This response is mediated in Arabidopsis , tomato and rice plants by a mobile protein known as FLOWERING LOCUS T (FT), the main component of the long-range florigen signal. A study of the potato ( Solanum tuberosum ) now shows that floral and tuberization transitions are controlled by two different FT -like genes ( StSP3D and StSP6A ) that respond to independent environmental cues. Seasonal fluctuations in day length regulate important aspects of plant development such as the flowering transition or, in potato ( Solanum tuberosum ), the formation of tubers. Day length is sensed by the leaves, which produce a mobile signal transported to the shoot apex or underground stems to induce a flowering transition or, respectively, a tuberization transition. Work in Arabidopsis, tomato and rice ( Oryza sativa ) identified the mobile FLOWERING LOCUS T (FT) protein as a main component of the long-range ‘florigen’, or flowering hormone, signal 1 , 2 , 3 . Here we show that expression of the Hd3a gene, the FT orthologue in rice, induces strict short-day potato types 4 to tuberize in long days. Tuber induction is graft transmissible and the Hd3a–GFP protein is detected in the stolons of grafted plants, transport of the fusion protein thus correlating with tuber formation. We provide evidence showing that the potato floral and tuberization transitions are controlled by two different FT -like paralogues (St SP3D and St SP6A ) that respond to independent environmental cues, and show that an autorelay mechanism involving CONSTANS modulates expression of the tuberization-control St SP6A gene.

α-Solanine and α-chaconine, steroidal glycoalkaloids (SGAs) found in potato (Solanum tuberosum), are among the best-known secondary metabolites in food crops. At low concentrations in potato tubers, SGAs are distasteful; however, at high concentrations, SGAs are harmful to humans and animals. Here, we show that POTATO GLYCOALKALOID BIOSYNTHESIS1 (PGA1) and PGA2, two genes that encode cytochrome P450 monooxygenases (CYP72A208 and CYP72A188), are involved in the SGA biosynthetic pathway, respectively. The knockdown plants of either PGA1 or PGA2 contained very little SGA, yet vegetative growth and tuber production were not affected. Analyzing metabolites that accumulated in the plants and produced by in vitro enzyme assays revealed that PGA1 and PGA2 catalyzed the 26- and 22-hydroxylation steps, respectively, in the SGA biosynthetic pathway. The PGA-knockdown plants had two unique phenotypic characteristics: The plants were sterile and tubers of these knockdown plants did not sprout during storage. Functional analyses of PGA1 and PGA2 have provided clues for controlling both potato glycoalkaloid biosynthesis and tuber sprouting, two traits that can significantly impact potato breeding and the industry.

• Light-induced tuber greening is one of the most important quality defects of potato. Although varietal and maturity factors are known to affect greening resistance, physiological mechanisms of resistance are poorly understood. We proposed that physiological and biochemical factors within the tuber periderm provide resistance and hypothesised that resistance is primarily related to suberin content. • We investigated differences in the tuber periderm between genotypes and tuber maturities that varied in greening propensity. We examined suberin and light-induced pigment accumulation, and phellem cell development and studied greening propensity in mutant and chemically treated tubers with enhanced suberisation. • Resistance to greening was strongly linked to increased suberin in the periderm, which varied with variety and tuber maturity. Furthermore, greening was reduced in mutant and chemically treated tubers with enhanced suberisation. Increases in phellem cell layers and light-induced carotenoids and anthocyanins were identified as secondary resistance factors. • Our work represents the first physiological mechanism of varietal and tuber maturity resistance to greening, expanding the known functionality of suberin and providing for the first time a biomarker that will aid producers and breeders in selection and improvement of potato varieties for greening resistance.

The absorption of the essential element nitrogen by plants affects various aspects of plant physiological activity, including gene expression, metabolite content and growth. However, the molecular mechanism underlying the potato tuberization response to nitrogen remains unclear. Potato plants were subjected to pot experiments under nitrogen deficiency, normal nitrogen levels and nitrogen sufficiency. A comprehensive analysis of the physiological responses, transcriptomic profiles, and metabolic pathways of potato stolons subjected to nitrogen stress was conducted. Transcriptomic analysis revealed 2756 differentially expressed genes (DEGs) associated with nitrogen stress. Metabolomic analysis identified a total of 600 differentially accumulated metabolites (DAMs). Further correlation analysis of the major DEGs and DAMs revealed that 9 key DEGs were associated with alpha-linolenic acid metabolism, 16 key DEGs with starch and sucrose metabolism, 7 key DEGs with nitrogen metabolism, and 16 key DEGs with ABC transporters. Nitrogen deficiency significantly increased the sucrose, GDP-glucose and L-glutamic acid levels and promoted stolon growth by increasing the expression of AMY (alpha-amylase), BE (1,4-alpha-glucan branching enzyme), SS (starch synthase), SPS (sucrose‒phosphate synthase) and AGPS (glucose‒1-phosphate adenylyltransferase). However, high nitrogen levels had the opposite effect. In addition, high nitrogen levels upregulated EG (endoglucanase), SUS (sucrose synthase) and GDH (glutamate dehydrogenase) and led to significant accumulation of 9-Hydroperoxy-10,12,15-octadecatrienoate (9(S)-HpOTrE), (13 S)-Hydroperoxyoctadeca-9,11,15-trienoate (13 (S)-HpOTrE) and L-glutamine, ultimately affecting the balance between plant growth and defense. Overall, our comprehensive study revealed the co-expressed genes and potential pathways related to potato tuber formation under different nitrogen conditions. These data provide a better understanding needed for improving potato tuber traits at the molecular and metabolic levels.

Potato (Solanum tuberosum) is the third most important food crop in the world. Potato tubers must be stored at cold temperatures to prevent sprouting, minimize disease losses, and supply consumers and the processing industry with high-quality tubers throughout the year. Unfortunately, cold storage triggers an accumulation of reducing sugars in tubers. High-temperature processing of these tubers results in dark-colored, bitter-tasting products. Such products also have elevated amounts of acrylamide, a neurotoxin and potential carcinogen. We demonstrate that silencing the potato vacuolar acid invertase gene VInv prevents reducing sugar accumulation in cold-stored tubers. Potato chips processed from VInv silencing lines showed a 15-fold acrylamide reduction and were light in color even when tubers were stored at 4°C. Comparable, low levels of VInv gene expression were observed in cold-stored tubers from wild potato germplasm stocks that are resistant to cold-induced sweetening. Thus, both processing quality and acrylamide problems in potato can be controlled effectively by suppression of the VInv gene through biotechnology or targeted breeding.

High temperature (HT) is a major environmental stress that severely inhibits potato (Solanum tuberosum L.) tuberisation and yield. Heat shock transcription factors (Hsfs) are pivotal in plant thermotolerance, yet their roles in potato remain unclear. Here, we demonstrate that overexpression of StHsfA2, a rapidly HT–responsive HSF family member, enhances thermotolerance and mitigates yield loss in transgenic potato under HT conditions. We reveal that StHsfA2 upregulates StSP6A expression by binding to the heat shock element–like motifs in its promoter. StSP6A encodes a homologue of FLOWERING LOCUS T that is critical for initiating tuber formation. Intriguingly, we found that StHsfA2 physically interacts with the StSP6A protein, which in turn inhibits StHsfA2–mediated StSP6A upregulation. However, HT stress attenuates the StHsfA2–StSP6A interaction. Thus, a negative feedback loop modulates StSP6A regulation by StHsfA2 under HT. In summary, our study shows that StHsfA2 is a key regulator of thermotolerance in potato plants. Its overexpression enhances heat resistance and could boost tuber yield, making it a promising candidate gene for countering yield loss amid global warming.