Julius-von-Sachs-Institut für Biowissenschaften
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- ACL (1)
- Ackerschmalwand (1)
- Aliphatics (1)
- Chain-length distribution (1)
- Cuticle (1)
- Cuticular water permeabilities (1)
- Cuticular waxes (1)
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- Homöostase (1)
The plant cuticle is a continuous extracellular protective layer covering the outermost surfaces of higher plants that are in contact with the surrounding atmosphere. The primary function of the cuticular lipid membrane, which is mainly composed of biopolymer cutin and cuticular waxes, is to protect the plant organs against uncontrolled water loss. The chemical composition and the biophysical properties of cuticular waxes affect the rate of water diffusion across the cuticle. Fruit transpiration plays an important role in the development and the maintenance of fruit quality. The fruit has been suggested to present better dehydration stress tolerance than the leaf. However, the differences in transpiration and the chemical composition of cuticular waxes between fruit and leaf have yet to be comprehensively investigated.
The present study aims to investigate the water permeability and cuticular wax composition of fruit and leaf cuticles of a wide range of plant species and to elucidate the different roles of the cuticular wax components in the transpiration barrier. To address these objectives, fruit and leaf samples from 17 species were investigated. The cuticular transpiration of intact fruits and astomatous adaxial leaf surfaces and the minimum leaf conductance obtained by leaf drying curves for intact leaves were gravimetrically determined for a variety of plant species. The chemical composition of cuticular waxes of fruits and leaves was thoroughly analysed by gas chromatography with flame ionization and mass spectrometry.
The water permeability of fruits ranged from 3.7 x 10-5 m s-1 (Prunus domestica subsp. syriaca) to 37.4 x 10-5 m s-1 (Coffea arabica), whereas permeability for leaves varied between 1.6 x 10-5 m s-1 (Cornus officinalis) and 4.5 x 10-5 m s-1 (Prunus domestica subsp. syriaca (L.)). The interspecies range of water permeability of fruits was significantly higher than that of leaves. Chemical analyses of the cuticular waxes demonstrated that fatty acids, primary alcohols, n-alkanes, aldehydes and alkyl esters were the predominant very-long-chain aliphatic compound classes of fruit and leaf surfaces. Sterols, such as β-sitosterol and campesterol, and triterpenoids, such as oleanolic acid, ursolic acid, α-amyrin and ß-amyrin, were the major cyclic compound classes in the cuticular wax membrane.
The amount and composition of cuticular waxes of both fruits and leaves varied at an intraspecific level. There were no significant correlations between the total cuticular wax load or the individual cuticular wax composition and the water permeability of fruits or leaves independently or together. After combining the fruit and leaf data set, a significant correlation between the average chain length of very-long-chain aliphatic compounds and permeabilities was detected, i.e. the longer the average chain length, the lower the water permeability.
Interestingly, n-Nonacosane (C29) was abundantly detected in fruit waxes of Rosaceae species. These fruits exhibited a relatively low transpiration level, which was very close to their leaf cuticular permeability. The present study suggests that the lower cuticular permeability of leaves, in comparison to that of fruits, may be attributed to the longer average chain length of aliphatic compounds. The accumulation of total wax, triterpenoids and aliphatic compounds may not contribute to the transpiration barrier directly. The present results are highly consistent with the previous model assumptions for the cuticular structure and transport barrier. Furthermore, this comparative study on leaf and fruit cuticles provides further insights linking the cuticular wax chemistry to the physiological properties of the plant cuticle.
The control of energy homeostasis is of pivotal importance for all living organisms. In the last years emerged the idea that many stress responses that are apparently unrelated, are actually united by a common increase of the cellular energy demand. Therefore, the so called energy signaling is activated by many kind of stresses and is responsible for the activation of the general stress response. In Arabidopsis thaliana the protein family SnF1- related protein kinases (SnRK1) is involved in the regulation of many physiological processes but is more known for its involvement in the regulation of the energy homeostasis in response to various stresses. To the SnRK1 protein family belong SnRK1.1 (also known as KIN10), SnRK1.2 (KIN11), and SnRK1.3 (KIN12). SnRK1 exerts its function regulating directly the activity of metabolic enzymes or those of key transcription factors (TFs). The only TFs regulated by SnRK1 identified so far is the basic leucine zipper (bZIP) 63. bZIP63 belongs to the C group of bZIPs (C-bZIPs) protein family together with bZIP9, bZIP10, and bZIP25. SnRK1.1 phosphorylates bZIP63 on three amino acids residues, serine (S) 29, S294, and S300. The phosphorylation of tbZIP63 is strongly related to the energy status of the plant, shifting from almost absent during the normal growth to strongly phosphorylated when the plant is exposed to extended dark. bZIPs normally bind the DNA as dimer in order to regulate the expression of their target genes. C-bZIPs preferentially form dimers with S1-bZIPs, constituting the so called C/S1- bZIPs network. The SnRk1 dependent phosphorylation of bZIP63 regulates its activation potential and its dimerization properties. In particular bZIP63 shift its dimerization preferences according to its phosphorylation status. The non-phosphorylated form of bZIP63 dimerize bZIP1, the phosphorylates ones, instead, forms dimer with bZIP1, bZIP11, and bZIP63 its self. Together with bZIP63, S1-bZIPs are important mediator of part of the huge transcriptional reprogramming induced by SnRK1 in response to extended dark. S1-bZIPs regulate, indeed, the expression of 4'000 of the 10'000 SnRK1-regulated genes in response to energy deprivation. In particular S1-bZIPs are very important for the regulation of many genes encoding for enzymes involved in the amino acid metabolism and for their use as alternative energy source. After the exposition for some hours to extended dark, indeed, the plant make use of every energy substrate and amino acids are considered an important energy source together with lipids and proteins. Interestingly, S1- bZIPs regulate the expression of ETFQO. ETFQO is a unique protein that convoglia the electrons provenienti from the branch chain amino acids catabolism into the mitochondrial electron transport chain. The dimer formed between bZIP63 and bZIP2 recruits SnRK1.1 directly on the chromatin of ETFQO promoter. The recruitment of SnRK1 on ETFQO promoter is associated with its acetylation on the lysine 14 of the histone protein 3 (K14H3). This chromatin modification is normally asociated with an euchromatic status of the DNA and therefore with its transcriptional activation. Beside the particular case of the regulation of ETFQO gene, S1-bZIPs are involved in the regulation of many other genes activated in response of different stresses. bZIP1 is for example an important mediator of the salt stress response. In particular bZIP1 regulates the primary C- and N-metabolism. The expression of bZIP1, in response of both salt ans energy stress seems to be regulated by SnRK1, as it is the expression of bZIP53 and bZIP63.
Beside its involvement in the regulation of the energy stress response and salt response, SnRK1 is the primary activators of the lipids metabolism during see germination. SnRK1, indeed, controls the expression of CALEOSINs and OLEOSINs. Those proteins are very important for lipids remobilization from oil droplets. Without their expression seed germination and subsequent establishment do not take place because of the absence of fuel to sustain these highly energy costly processes, which entirely depend on the catabolism of seed storages.