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2,4-D retards senescence of postharvest citrus fruits by increasing exogenous auxin, endogenous ABA and SA contents, while decreasing ethylene production; and enhancing stress-defense capability through changing epicuticular wax morphology and lignin content in peel

The use of plant-based indicators and other conventional means to detect the level of water stress in crops may be challenging, due to their difficulties in automation, their arduousness, and their time-consuming nature. Non-contact and non-destructive sensing methods can be used to detect the level of water stress in plants continuously and to provide automatic sensing and controls. This research aimed at determining the viability, efficiency, and swiftness in employing the commercial Workswell WIRIS Agro R infrared camera (WWARIC) in monitoring water stress and scheduling appropriate irrigation regimes in mandarin plants. The experiment used a four-by-three randomized complete block design with 80–100% FC water treatment as full field capacity and three deficit irrigation treatments at 70–75% FC, 60–65% FC, and 50–55% FC. Air temperature, canopy temperature, and vapor pressure deficits were measured and employed to deduce the empirical crop water stress index, using the Idso approach (CWSI(Idso)) as well as baseline equations to calculate non-water stress and water stressed conditions. The relative leaf water content (RLWC) of mandarin plants was also determined for the growing season. From the experiment, CWSI(Idso) and CWSI were estimated using the Workswell Wiris Agro R infrared camera (CWSIW) and showed a high correlation (R2 = 0.75 at p < 0.05) in assessing the extent of water stress in mandarin plants. The results also showed that at an altitude of 12 m above the mandarin canopy, the WWARIC was able to identify water stress using three modes (empirical, differential, and theoretical). The WWARIC’s color map feature, presented in real time, makes the camera a suitable device, as there is no need for complex computations or expert advice before determining the extent of the stress the crops are subjected to. The results prove that this novel use of the WWARIC demonstrated sufficient precision, swiftness, and intelligibility in the real-time detection of the mandarin water stress index and, accordingly, assisted in scheduling irrigation.

Background Boron (B)-toxicity is an important disorder in agricultural regions across the world. Seedlings of `Sour pummelo' ( Citrus grandis ) and `Xuegan' ( Citrus sinensis ) were fertigated every other day until drip with 10 μM (control) or 400 μM (B-toxic) H 3 BO 3 in a complete nutrient solution for 15 weeks. The aims of this study were to elucidate the adaptive mechanisms of citrus plants to B-toxicity and to identify B-tolerant genes. Results B-toxicity-induced changes in seedlings growth, leaf CO 2 assimilation, pigments, total soluble protein, malondialdehyde (MDA) and phosphorus were less pronounced in C. sinensis than in C. grandis . B concentration was higher in B-toxic C. sinensis leaves than in B-toxic C. grandis ones. Here we successfully used cDNA-AFLP to isolate 67 up-regulated and 65 down-regulated transcript-derived fragments (TDFs) from B-toxic C. grandis leaves, whilst only 31 up-regulated and 37 down-regulated TDFs from B-toxic C. sinensis ones, demonstrating that gene expression is less affected in B-toxic C. sinensis leaves than in B-toxic C. grandis ones. These differentially expressed TDFs were related to signal transduction, carbohydrate and energy metabolism, nucleic acid metabolism, protein and amino acid metabolism, lipid metabolism, cell wall and cytoskeleton modification, stress responses and cell transport. The higher B-tolerance of C. sinensis might be related to the findings that B-toxic C. sinensis leaves had higher expression levels of genes involved in photosynthesis, which might contribute to the higher photosyntheis and light utilization and less excess light energy, and in reactive oxygen species (ROS) scavenging compared to B-toxic C. grandis leaves, thus preventing them from photo-oxidative damage. In addition, B-toxicity-induced alteration in the expression levels of genes encoding inorganic pyrophosphatase 1, AT4G01850 and methionine synthase differed between the two species, which might play a role in the B-tolerance of C. sinensis . Conclusions C. sinensis leaves could tolerate higher level of B than C. grandis ones, thus improving the B-tolerance of C. sinensis plants. Our findings reveal some novel mechanisms on the tolerance of plants to B-toxicity at the gene expression level.

This work was supported by Ministerio de Economía y Competitividad (MINECO) and Universitat Jaume I through grants No. AGL2013-42038-R and P1IB2013-23, respectively. SIZ was supported by a predoctoral grant from Universitat Jaume I.

Key message This work reveals the protective role of two rhizobacteria, Pseudomonas putida and Novosphingobium sp., on citrus plants subjected to salt stress conditions. Abstract Detrimental salt stress effects on crops are likely to increase due to climate change reducing the quality of irrigation water. Plant growth-promoting rhizobacteria (PGPRs) can mitigate stress-induced damage in plants cultivated under high salinity conditions. In this work, Citrus macrophylla (alemow) plants inoculated with the rhizobacteria Pseudomonas putida KT2440 or Novosphingobium sp. HR1a were subjected to salt stress for 30 days. Results showed that in absence of salt stress, Novosphingobium sp. HR1a induced a decrease of transpiration (E) and stomatal conductance (gs). Both rhizobacteria reduced salt stress-induced damage. Levels of abscisic acid (ABA) and salicylic acid (SA) were lower in inoculated plants under salt stress conditions. Similarly, under stress conditions maximum efficiency of photosystem II (Fv/Fm) in inoculated plants decreased to a lower extent than in non-inoculated ones. In stressed plants, Novosphingobium sp. HR1a also induced leaf accumulation of 3-indole acetic acid (IAA) and a delay in the decrease of quantum yield (ΦPSII). P. putida KT2440 inhibited root chloride and proline accumulation in response to salt stress. Although both bacterial species had beneficial effects on salt-stressed citrus plants, Novosphingobium sp. HR1a induced a better plant performance. Therefore, both strains could be candidates to be used as PGPRs in programs of inoculation for citrus protection against salt stress.

Plant defense compounds occur in floral nectar, but their ecological role is not well understood. We provide evidence that plant compounds pharmacologically alter pollinator behavior by enhancing their memory of reward. Honeybees rewarded with caffeine, which occurs naturally in nectar of Coffea and Citrus species, were three times as likely to remember a learned floral scent as were honeybees rewarded with sucrose alone. Caffeine potentiated responses of mushroom body neurons involved in olfactory learning and memory by acting as an adenosine receptor antagonist. Caffeine concentrations in nectar did not exceed the bees' bitter taste threshold, implying that pollinators impose selection for nectar that is pharmacologically active but not repellent. By using a drug to enhance memories of reward, plants secure pollinator fidelity and improve reproductive success.

Asian citrus psyllid (Diaphorina citri) feeding behaviors play a significant role in the transmission of the phloem-limited Candidatus Liberibacter asiaticus (CLas) bacterium that causes the economically devastating citrus greening disease. Sustained phloem ingestion by D. citri on CLas infected plants is required for pathogen acquisition and transmission. Recent studies have shown a fibrous ring of thick-walled sclerenchyma around the phloem in mature, fully expanded citrus leaves that is more prominent on the abaxial compared with the adaxial side. The composition and thickness of this fibrous ring may have an important role in selection of feeding sites by D. citri based on leaf age and leaf surface, which in turn can affect pathogen acquisition and transmission. We measured feeding behavior using electrical penetration graph (EPG) recordings of individual D. citri adults placed on abaxial or adaxial surfaces of young or mature Valencia orange leaves to study the role of the sclerenchymatous ring in modifying D. citri feeding behavior. Feeding sites on the same leaf tissues were then sectioned and examined by epifluorescence microscopy. The duration of phloem ingestion (E2 waveform) by psyllids was significantly reduced on mature compared with young leaves, and on abaxial compared with adaxial leaf surfaces. The longest duration of phloem ingestion was observed from psyllids placed on the adaxial side of young leaves that had the least developed sclerenchyma. Bouts of phloem salivation (E1 waveform), however, were significantly longer on mature leaves compared with young leaves. D. citri adults made consecutive phloem feeding attempts (bouts) on the abaxial side of mature leaves and those bouts resulted in unsuccessful or shorter periods of phloem ingestion. Adults also made more frequent and longer bouts of xylem ingestion on mature leaves compared with adult psyllids placed on young leaves. Epifluorescence microscopy showed that the fibrous ring in young leaves was thinner and autofluoresced in red whereas the ring in mature leaves was thicker and autofluoresced in blue, indicating changes in structure and composition (e.g., lignification) of sclerenchyma correlated with leaf age. Our results support the hypothesis that the presence of a thick, well-developed fibrous ring around phloem tissues of mature leaves acts as a barrier to frequent or prolonged phloem ingestion by D. citri from citrus leaves. This may have an important role in limiting or preventing CLas acquisition and/or transmission by D. citri, and could be used for identification and development of resistant citrus cultivars.

The long juvenile period of citrus trees (often more than 6 years) has hindered genetic improvement by traditional breeding methods and genetic studies. In this work, we have developed a biotechnology tool to promote transition from the vegetative to the reproductive phase in juvenile citrus plants by expression of the Arabidopsis thaliana or citrus FLOWERING LOCUS T (FT) genes using a Citrus leaf blotch virus-based vector (clbvINpr-AtFT and clbvINpr-CiFT, respectively). Citrus plants of different genotypes graft inoculated with either of these vectors started flowering within 4–6 months, with no alteration of the plant architecture, leaf, flower or fruit morphology in comparison with noninoculated adult plants. The vector did not integrate in or recombine with the plant genome nor was it pollen or vector transmissible, albeit seed transmission at low rate was detected. The clbvINpr-AtFT is very stable, and flowering was observed over a period of at least 5 years. Precocious flowering of juvenile citrus plants after vector infection provides a helpful and safe tool to dramatically speed up genetic studies and breeding programmes.

Citrus Huanglongbing (HLB), a highly devastating citrus disease, is associated with 'Candidatus Liberibacter asiacitus' (CLas), a member of phloem-inhabiting α-proteobacteria. HLB can affect all cultivated citrus and no cure is currently available. Previous studies showed that Kaffir lime (Citrus hystrix), primarily grown in South Asia and Southeast Asia, was tolerant to HLB but the molecular mechanism remains unknown. In this study, gene expression profiling experiments were performed on HLB-tolerant C. hystrix and HLB-susceptible C. sinensis three months after inoculation with CLas using RNA-seq data. Differentially expressed genes (DEGs) in the two citrus cultivars were mainly involved in diverse cellular functions including carbohydrate metabolism, photosynthesis, cell wall metabolism, secondary metabolism, hormone metabolism and oxidation/reduction processes. Notably, starch synthesis and photosynthesis process were not disturbed in CLas-infected C. hystrix. Most of the DEGs involved in cell wall metabolism and secondary metabolism were up-regulated in C. hystrix. In addition, the activation of peroxidases, Cu/Zn-SOD and POD4, may also enhance the tolerance of C. hystrix to CLas. This study provides an insight into the host response of HLB-tolerant citrus cultivar to CLas. C. hystrix is potentially useful for HLB-tolerant/resistant citrus breeding in the future.

Rapid assessment of physiological status is a precondition for addressing biological stress in trees so that they may recover. Environmental stress can cause water deficit in plants, while terahertz (THz) spectroscopy is sensitive to changes in aqueous solutions within organisms. This has given the THz sensor a competitive edge for evaluating plant phenotypes, especially under similar environmental stress, if there are existing differences in the corresponding THz information. In this study, we utilized THz technology in association with traditional weighing methods to explore physiological changes in citrus leaves under different temperature, duration, and stress treatment conditions. It was found that the higher the temperature and the longer the exposure duration, the more severe the reduction in the relative absorption coefficient. There was a positive correlation between the trends and the increase in the ion permeability of cells. In addition, based on the effective medium theory, THz spectral information can be transformed into information on free water and bound water in the leaves. Under different treatment conditions, water content shows different trends and degrees of change on the time scale, and accuracy was verified by traditional weighing methods. These findings revealed that characteristics of THz information can serve as a simple and clear indicator for judging a plant’s physiological status.