POSTHARVEST BIOLOGY AND TECHNOLOGY

From laboratory to industrial storage - Translating volatile organic compounds into markers for assessing garlic storage quality
Ludlow RA, Evans G, Graz M, Marti G, Martínez PC, Rogers HJ and Müller CT
Garlic ( L.) has long been grown for its culinary and health-promoting qualities. The seasonal nature of garlic cropping requires that bulbs be stored for many months after harvest to ensure a year-round supply. During this time, quality is known to deteriorate, and efforts have been made to improve the longevity of stored bulbs. Cold temperatures within the stores prolong shelf life, but fine temperature control is needed to avoid freezing damage or cold induced stress. Here, quality traits (alliinase activity, firmness, and water content) are measured in response to a 96 h - 5 °C cold stress, to simulate the effect of non-isothermic temperature control in a - 1.5 °C warehouse. Volatile organic compounds (VOCs) are measured by thermal desorption gas chromatography time of flight mass spectrometry to identify markers of non-isothermic storage in garlic. 129 compounds were putatively identified and four (L-lactic acid, 2,6-dimethylhetpadecane, 4-methyldodecane, and methylcyclopentane) showed high predictive accuracy for cold stress. VOCs were also sampled directly from a cold storage facility and the whole profile discriminated between sampling time points. Five VOCS were highly predictive for storage time in the warehouse but were different to VOCs previously shown to discriminate between storage times in a laboratory setting. This indicates the need for realistic warehouse experiments to test quality markers.
Investigating the role of abscisic acid and its catabolites on senescence processes in green asparagus under controlled atmosphere (CA) storage regimes
Anastasiadi M, Collings ER and Terry LA
Asparagus () is a highly perishable crop with a short postharvest life. Although some research has been done on the application of controlled atmosphere (CA), it has not been sufficiently explored and the underlying mechanisms controlling asparagus senescence processes are not well understood, restricting its potential for commercial application. The aim of this study was to investigate for the first time the link between abscisic acid (ABA) and ABA catabolites and senescence in asparagus stored under a range of different CA conditions. Two different set-ups were run in parallel; a traditional CA delivered by an International Controlled Atmosphere (ICA) system with continuous gas supply and LabPods™ fitted with sensors for real time monitoring of respiration rate (RR) and respiratory quotient (RQ) and able to retain established CA conditions with minimum gas supply requirements. The role of genetic variability was also studied by including two UK grown asparagus cultivars 'Gijnlim' and 'Jaleo' adapted for different climatic conditions. The results indicated that ABA and its catabolites were present in significantly higher concentrations in the air stored spears (control) compared to CA throughout storage, irrespective of cultivar, and were associated with accelerated senescence processes observed in control samples, such as textural changes indicative of spear toughening, discolouration, sugar depletion and asparagine accumulation. Furthermore, partial least squares regression (pls-r) applied for both cultivars, successfully differentiated samples based on O and CO concentrations and storage duration, both in cold storage and during shelf-life with the separation being driven primarily by ABA and its catabolites. Physiological and biochemical results indicated that all three CA conditions tested ([CA1] 2.5% O, 3% CO, [CA2] 2.5% O, 6% CO and [CA3] 2.5% O, 10% CO) successfully retained quality parameters including texture, colour, moisture content and visual appearance longer compared to air (control); however, they did not completely suppress the development of 'tip-breakdown' (a physiological disorder also known as tip rot) towards the end of storage, which coincided with rising concentrations of phaseic acid indicating an activation of the abscisic biosynthetic and catabolic pathway. It can be concluded that CA conditions can delay senescence for at least 3-weeks (2 weeks cold storage and 1 week shelf-life), by lowering metabolic rate and respiratory quotient (RQ) within the spears compared to control, and through successfully regulating ABA biosynthetic and catabolic pathways.
New insights into the effects of ethylene on ABA catabolism, sweetening and dormancy in stored potato tubers
Tosetti R, Waters A, Chope GA, Cools K, Alamar MC, McWilliam S, Thompson AJ and Terry LA
Continuous ethylene supplementation suppresses postharvest sprouting, but it can increase reducing sugars, limiting its use as an alternative to chlorpropham for processing potatoes. To elucidate the mechanisms involved, tubers were treated after curing with or without the ethylene binding inhibitor 1-methylcyclopropene (1-MCP at 1 μL L for 24 h), and then stored in air or air supplemented with continuous ethylene (10 μL L). Across three consecutive seasons, changes in tuber physiology were assessed alongside transcriptomic and metabolomic analysis. Exogenous ethylene alone consistently induced a respiratory rise and the accumulation of undesirable reducing sugars. The transient respiratory peak was preceded by the strong upregulation of two genes encoding 1-aminocyclopropane-1-carboxylate oxidase (ACO), typical of wound and stress induced ethylene production. Profiles of parenchymatic tissue highlighted that ethylene triggered abscisic acid (ABA) catabolism, evidenced by a steep fall in ABA levels and a transient rise in the catabolite phaseic acid, accompanied by upregulation of transcripts encoding an ABA 8'-hydroxylase. Moreover, analysis of non-structural carbohydrate-related genes revealed that ethylene strongly downregulated the expression of the , already known to be involved in cold-induced sweetening. All these ethylene-induced effects were negated by 1-MCP with one notable exception: 1-MCP enhanced the sprout suppressing effect of ethylene whilst preventing ethylene-induced sweetening. This study supports the conclusions that: i) tubers adapt to ethylene by regulating conserved pathways ( ABA catabolism); ii) ethylene-induced sweetening acts independently from sprout suppression, and is similar to cold-induced sugar accumulation.
Innovative breeding technologies in lettuce for improved post-harvest quality
Damerum A, Chapman MA and Taylor G
Societal awareness of healthy eating is increasing alongside the market for processed bagged salads, which remain as one of the strongest growing food sectors internationally, including most recently from indoor growing systems. Lettuce represents a significant proportion of this ready-to-eat salad market. However, such products typically have a short shelf life, with decay of post-harvest quality occurring through complex biochemical and physiological changes in leaves and resulting in spoilage, food waste and risks to health. We review the functional and quantitative genetic understanding of lettuce post-harvest quality, revealing that few findings have translated into improved cultivar development. We identify (i) phytonutrient status (for enhanced antioxidant and vitamin status, aroma and flavour) (ii) leaf biophysical, cell wall and water relations traits (for longer shelf life) (iii) leaf surface traits (for enhanced food safety and reduced spoilage) and (iv) chlorophyll, other pigments and developmental senescence traits (for appearance and colour), as key targets for future post-harvest breeding. Lettuce is well-placed for rapid future exploitation to address postharvest quality traits with extensive genomic resources including the recent release of the lettuce genome and the development of innovative breeding technologies. Although technologies such as CRISPR/Cas genome editing are paving the way for accelerated crop improvement, other equally important resources available for lettuce include extensive germplasm collections, bi-parental mapping and wide populations with genotyping for genomic selection strategies and extensive multiomic datasets for candidate gene discovery. We discuss current progress towards post-harvest quality breeding for lettuce and how such resources may be utilised for future crop improvement.
Transcriptome and phytohormone changes associated with ethylene-induced onion bulb dormancy
Alamar MC, Anastasiadi M, Lopez-Cobollo R, Bennett MH, Thompson AJ, Turnbull CGN, Mohareb F and Terry LA
Control of dormancy and sprouting in onion bulbs is commercially important for postharvest management. Although ethylene application is sometimes used to extend dormancy, the underlying mechanisms regulating dormancy transition remain unclear. Since the sprout leaves emerge from the bulb baseplate, we used this tissue to assess the impact of ethylene treatment and storage time on the hormone profile and the transcriptome. Reads from 30 libraries were assembled and annotated, with 94,840 unigenes retained after filtering. The transcriptome assembly was of high quality and continuity (N50: 1809 bp, GC content: 36.21 %), and was used to analyse differential expression and Gene Onotologies. Across two years, applied ethylene resulted in delayed dormancy break and reduced post-dormancy sprout vigour. Ethylene supplementation enhanced endogenous ethylene production and caused a transient climacteric-like increase in respiration. Significant changes in hormone and associated transcript profiles occurred through storage and in response to ethylene. In particular, abscisic acid (ABA) and its metabolite phaseic acid (PA) increased under ethylene during the longer dormancy period; however, cytokinin increases observed during storage appeared largely independent of ethylene treatment. Several hormone-related transcripts showed differential expression over time and/or in response to ethylene. Expression of ethylene biosynthesis (), receptor () and transcription factor () genes were modified by ethylene, as were ABA biosynthesis genes such and cytokinin biosynthesis genes such as and . We conclude that ethylene substantially modifies expression of genes in several phytohormone pathways, and some of these changes may underlie the dormancy-extending effects of exogenous ethylene.
Growth temperature influences postharvest glucosinolate concentrations and hydrolysis product formation in first and second cuts of rocket salad
Jasper J, Wagstaff C and Bell L
Rocket salad species ( and ; also known as ) are known for their high concentrations of health-related isothiocyanates, which are derived from secondary metabolites called glucosinolates. Increases in temperature due to climate change and extreme weather event frequencies over the coming decades are likely to influence not only the growth of leafy vegetables, but also their nutritional density. It is therefore essential to determine the impacts of these in order to mitigate crop losses and nutritional decline in future. Our data show there is a strong influence of pre-harvest growth temperatures on glucosinolate biosynthesis and formation of glucosinolate hydrolysis products postharvest, and that this is genotype dependent. High growth temperature (40 °C) severely retarded germination, growth, regrowth, and survival of rocket plants. Highest glucosinolate concentrations were observed in first and second cuts at 40 °C, but did not correspond to highest isothiocyanate concentrations (observed at 30 °C, second cut). Hydrolysis product formation is proportionately not as great as glucosinolate increases at 40 °C, possibly due to inhibition of enzyme function(s) at higher temperatures. These data indicate that high growth temperatures increase glucosinolate accumulation, but growth and productivity is significantly reduced. Much greater emphasis is needed for breeding cultivars tolerant to high growth temperatures in order to maximise nutritional benefits imparted by temperature stress.
Seasonal and temporal changes during storage affect quality attributes of green asparagus
Anastasiadi M, Collings ER, Shivembe A, Qian B and Terry LA
Asparagus is a perennial crop with a short UK harvest season. Methods to extend the storage life of asparagus have proven difficult. To gain insight into the physiological (. colour, respiration rate, cutting energy, and stiffness measured using laser Doppler vibrometry), and biochemical (. sugars, ascorbic acid, and abscisic acid and its catabolites) changes throughout the UK season, two cultivars were harvested weekly and stored under shelf life conditions (7 °C). Results were compared to spears (plus one additional cultivar) cold stored (1 °C) for three weeks followed by one week of shelf life. Concentrations of sugar, abscisic acid (ABA) and catabolites at harvest were subject to seasonal variation, directly affecting storage potential. A generalised linear model with stepwise feature selection was applied to select the most important parameters for the prediction of total sugars and phaseic acid (PA). More favourable growing conditions at harvest increased sugars and lowered ABA content and catabolites, which coincided with better maintenance of spear quality during storage; including maintaining textural characteristics. Storage time had a negative impact on spear texture and sugar content, with cutting energy increasing and stiffness decreasing both during cold storage and subsequent shelf life. A partial shift in sugar biosynthesis occurred during shelf life increasing sucrose concentrations. Results suggest that the temporal flux in ABA and catabolites, and individual sugars could be used to model storage potential of asparagus spears.
Fructans redistribution prior to sprouting in stored onion bulbs is a potential marker for dormancy break
Ohanenye IC, Alamar MC, Thompson AJ and Terry LA
Continuous supply of high quality onion bulbs to meet year-round demand is dependent on maintaining dormancy and bulb quality during storage. Sprouting impacts negatively on the storage quality of onion bulbs. Ethylene supplementation has previously been revealed to inhibit sprout growth in stored onion bulbs. Fructans content, especially those at higher degree of polymerisation (DP), are reported to positively correlate with delayed sprouting. However, little is known about the impact of pre-harvest irrigation regimes on fructans accumulation and redistribution in relation to onion bulb dormancy and quality in store. Across two seasons, onion plants of cultivars 'Red Baron' and 'Sherpa' were subjected to full irrigation (FI) (100% replenishment of crop evapotranspiration) or deficit irrigation (DI) (50% of FI treatment) from bulb initiation to harvest. Bulbs were harvested at full maturity and stored at 1 °C for five months. Bulbs were treated with or without 1-MCP (1 μL L) for 24 h before storage under continuous ethylene supplementation (10 μL L) or air. DI had no effect on dormancy-break, sprout emergence, total fructans content and total sugar content. In contrast, ethylene delayed sprout emergence and suppressed sprout growth; added 1-MCP enhanced this effect. The concentration of DP3-8 fructans were higher in top and bottom sections compared to the baseplate. Before sprout emergence, fructans of DPs 7-8 were no longer present in the top and bottom wedges, while they accumulated in the baseplate; irrespective of pre- or postharvest treatments. This redistribution of fructans within the bulb suggested a transition in dormancy state and could be used as a predictive marker for sprouting in stored onion bulbs.
Effect of UV-C on the physiology and biochemical profile of fresh berries
Collings ER, Alamar Gavidia MC, Cools K, Redfern S and Terry LA
Application of UV-C has been shown to enhance the biochemical profile of various plant materials. This could be used to increase biochemical load, reducing the amount of material required but still impart equivalent flavour. As spices, such as black pepper ( L.), are typically dried to low moisture content to create a stable product for transportation and storage, little work has explored the use of modern postharvest treatments to enhance flavour. In this work, fresh berries were exposed to four UV-C doses (0, 1, 5 and 15 kJ m) and subsequently stored at 5 °C for . 4 weeks. Two separate experiments (early and late season) were conducted across one season. Replicate berry clusters were stored separately within continuously ventilated 13 L boxes. Real-time respiration rate (), ethylene production, fruit colour and water potential were measured at regular intervals during storage. In addition, piperine and essential oils were assessed using a simple newly developed method which enabled both compound groups to be simultaneously extracted and subsequently quantified. UV-C was found to cause significant changes in colour (from green to brown) whilst also altering the biochemical composition (piperine and essential oils), which was influenced by UV-C dose and berry maturity. Low to medium UV-C doses could potentially enhance flavour compounds in black pepper enabling processors to create products with higher biochemical load.
Tissue biochemical diversity of 20 gooseberry cultivars and the effect of ethylene supplementation on postharvest life
Anastasiadi M, Mwangi PM, Ordaz-Ortiz JJ, Redfern SP, Berry M, Simmonds MSJ and Terry LA
The European gooseberry () is still an understudied crop with limited data available on its biochemical profile and postharvest life. A variety of polyphenols were detected in the skin and flesh of 20 gooseberry cvs, representing mainly flavonol glycosides, anthocyanins and flavan-3-ols. In contrast, gooseberry seeds were for the first time characterised by the presence of considerable amounts of hydroxycinnamic acid glycosides tentatively identified by UPLC-QToF/MS. All cvs examined represented a good source of vitamin C while being low in sugar. Furthermore, the postharvest stability of bioactives was explored by supplementation of exogenous ethylene in air at 5 °C. Results suggest a low sensitivity of gooseberries to ethylene. The overall quality of gooseberries remained stable over two weeks, showing potential for extended bioactive life.
Heat treatment prevents postharvest geotropic curvature of asparagus spears (Asparagus officinalis L.)
Paull RE and Chen NJ
Negative geotropic curvature of fresh asparagus spears after harvest was prevented by a brief heat treatment. The heat treatment was immersion in heated water at 47.5 degrees C for 2-5 min and cooling to storage temperature as soon as possible after heat treatment. The heat treatment can be varied from 45 degrees C for longer than 5 min to 50 degrees C for 2.5 min or less to avoid significant loss of spear appearance. The treatment temperature and time needed to be adjusted for spear diameter, small diameter spears required a shorter exposure time or lower temperature.