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Infrared and Raman Spectroscopic Features of Plant Cuticles: a Review

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Infrared and Raman Spectroscopic Features of Plant Cuticles: a Review REVIEW ARTICLE published: 25 June 2014 doi: 10.3389/fpls.2014.00305 Infrared and Raman spectroscopic features of plant cuticles: a review José A. Heredia-Guerrero 1*, José J. Benítez 2, Eva Domínguez 3, Ilker S. Bayer 1, Roberto Cingolani 4, Athanassia Athanassiou 1 and Antonio Heredia 3,5 1 Nanophysics, Istituto Italiano di Tecnologia, Genova, Italy 2 Instituto de Ciencias de Materiales de Sevilla, CSIC-US, Seville, Spain 3 Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora, CSIC-UMA, Málaga, Spain 4 Istituto Italiano di Tecnologia, Genova, Italy 5 Departamento de Bioquímica y Biología Molecular, Facultad de Ciencias, Universidad de Málaga, Málaga, Spain Edited by: The cuticle is one of the most important plant barriers. It is an external and continuous Andreia Michelle Smith-Moritz, lipid membrane that covers the surface of epidermal cells and whose main function is to Lawrence Berkeley Labs, USA prevent the massive loss of water. The spectroscopic characterization of the plant cuticle Reviewed by: and its components (cutin, cutan, waxes, polysaccharides and phenolics) by infrared Barbara G. Pickard, Washington University in St. Louis, USA and Raman spectroscopies has provided significant advances in the knowledge of the Thomas Eichert, University of Bonn, functional groups present in the cuticular matrix and on their structural role, interaction Germany and macromolecular arrangement. Additionally, these spectroscopies have been used *Correspondence: in the study of cuticle interaction with exogenous molecules, degradation, distribution José A. Heredia-Guerrero, Smart of components within the cuticle matrix, changes during growth and development and Materials Group, Nanophysics, Istituto Italiano di Tecnologia, Via characterization of fossil plants. Morego 30, Genova, 16163, Italy e-mail: [email protected] Keywords: plant cuticle, cuticle components, cuticle structure, infrared spectroscopy, Raman spectroscopy If I am to know an object, though I need not know Cutan can partially or completely substitute cutin as the cuti- its external properties, I must know all its internal cle matrix. Significant differences in cuticle composition can be properties. observed among plants, different organs within a plant or even Ludwig Wittgenstein (Tractatus among developmental stages of a given organ. Similarly, environ- Logico-Philosophicus, 1922) mental conditions can modify the amount and composition of the cuticle (Domínguez et al., 2012). More details about the chemical INTRODUCTION composition and the spatial distribution of cuticle components The plant cuticle is the most external and continuous membrane can be found elsewhere (Jeffree, 2006; Pollard et al., 2008). that covers epidermal cells of leaves, fruits, petals, and non- The cuticle is one of the most important plant barriers. In this lignified stems (Heredia, 2003). It is a composite membrane with sense, the biophysical properties of plant cuticles, structural, ther- a heterogeneous spatial distribution, Figure 1. The matrix is com- mal, biomechanical, and hydric, are a complex balance between posed of cutin, a long-chain and insoluble polymer formed by their protective role and the necessity of the plant to grow and hydroxylated and epoxy-hydroxylated C16 and C18 esterified fatty develop (Domínguez et al., 2011).Themainfunctionascribedto acids. The inner side is rich in polysaccharides (cellulose, hemi- the cuticle is the protection of plants against uncontrolled water celluloses, and pectins) from the plant cell wall, and represents loss (Burghardt and Riederer, 2006). Additionally, as an interface the attachment site to the outer epidermal cell wall. Other cuticle between the plant and the environment, it has other secondary components are soluble waxes (mixtures of homologous series of roles (Yeats and Rose, 2013): it represents the first defense against long-chain aliphatics, such as alkanes, alcohols, aldehydes, fatty pests and pathogens, it can efficiently reflect dangerous UV light acids and esters, together with variable amounts of cyclic com- (depending on the crystallinity of the epicuticular waxes), and it is pounds such as triterpenoids) located on the surface (epicuticular involved in the establishment of organ boundaries during devel- waxes) or distributed through the cuticle (intracuticular waxes), opment. Plants with superhydrophobic cuticles (Lotus effect) and phenolic compounds such as cinnamic acids and flavonoids. have further biological advantages in terms of self-cleaning and Cuticles from some species may contain an alternative, and also reduction of water content in the surface. The self-cleaning of chemically inert, polymer known as cutan, which is thought to cuticles can provide an additional defense against the deposition consist of an ether-linked network of methylene chains, double of pathogens and sunlight-blocking particles, while the reduction bonds, and carboxyl groups (Villena et al., 1999; Jeffree, 2006). of water can slow down the growth of microorganism and the leaching of nutrients (Koch and Barthlott, 2009; Yeats and Rose, Abbreviations: FTIR, Fourier Transform Infrared; ATR, Attenuated Total 2013). Reflection; DRIFT, Diffuse Reflectance Infrared Fourier Transform; NIR, Near Infrared; CARS, Coherent Anti-Stokes Raman Spectroscopy; TIR, Total Internal In addition to the biological and agricultural importance Reflection; DMSO, dimethyl sulfoxide. of this plant barrier, its applied use as a source of organic www.frontiersin.org June 2014 | Volume 5 | Article 305 | 1 Heredia-Guerrero et al. IR/Raman features of plant cuticles acid/carboxylate functional groups), and non-degradable frac- tions cannot be analyzed (Pollard et al., 2008). Additionally, they have substantial weaknesses regarding the structural determina- tion of such components. More traditional structural techniques as X-ray diffraction have scarcely been used due to the amor- phous nature of the plant cuticle (Luque et al., 1995). In contrast, solid state 13C nuclear magnetic resonance, using cross or direct polarization and magic-angle spinning methods, allows the iden- tification of functional groups and structures, the quantification of the molecular dynamics and the assessment of the cross-linking capability. Nonetheless, these spectroscopic measurements pro- vide limited information concerning molecular structure due to overlapping of the broad spectral lines and, for quantitative mea- surements, long acquisition times are required (Serra et al., 2012). IR and Raman spectroscopies are non-destructive and accessi- ble techniques which have shown important advantages in the chemical and structural analysis of plant cuticles, e.g., identifi- cation of functional groups and conformations, determination of intra- and intermolecular interactions of cuticle components with exogenous molecules, and qualitative measurements of the cutin polymerization. These spectroscopies are based on the excitation of the molecular vibrations of chemical bonds by the absorp- tion of light (infrared spectroscopy) or the inelastic scattering of photons (Raman spectroscopy). Both phenomena are gov- erned by different mechanisms, affecting the exact position, the FIGURE 1 | Schematic diagram of a transverse section of a plant appearance and intensity of the bands in the corresponding spec- cuticle. The figure shows the typical shape of the plant cuticle between two epidermal cells. Main components and their space distribution are tra. Main advantages of Raman spectroscopy are the possibility displayed: the layer below is rich in polysaccharides from the cell wall, while of using water as solvent and practically no sample prepara- the top layer is mainly constituted by cutin with a last layer of epicuticular tion. Nevertheless, this spectroscopy presents some drawbacks. waxes. Intracuticular waxes and phenolic compounds are spread through For example, fluorescence may interfere with the mechanism of the plant cuticle. the Raman effect and overlap the signals. On the other hand, IR measurements are fast, easy and no interferences are produced compounds and its importance as plant biomass have recently by other mechanisms. However, IR spectroscopy is very sensitive begun to be considered. In this regard, cuticle components are to water and it cannot be used as solvent. Also, the prepara- a potential alternative feedstock for aliphatic compounds com- tion of samples for IR spectroscopy presents some limitations monly found in oil plants (Tsubaki and Azuma, 2013). Thus, the on sample thickness, uniformity and dilution to avoid satura- potential applied value of tomato fruit peel, grape skins and green tion. Furthermore, the wide set of different modes of acquisition tea residues have been assessed (Arrieta-Baez et al., 2011; Mendes of infrared and Raman techniques can provide important and et al., 2013; Tsubaki and Azuma, 2013). Some of these aliphatic complementary chemical information. For instance, in the trans- compounds, specifically cutin monomers, have been used to syn- mission mode the sample can be placed directly into the path thetize long-chain polyesters, resulting in polymers with similar of the infrared beam, providing information about whole sys- characteristics to the plant cutin (Benítez et al., 2004a; Heredia- tem. Other mode of acquisition is ATR-FTIR. ATR is a useful Guerrero et al., 2009; Gómez-Patiño et al., 2013; Vilela et al., technique to obtain the IR spectrum of the surface of samples. 2014). The sample is placed in contact with an internal reflection ele- The lipid
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