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Plant Latex, from Ecological Interests to Bioactive Chemical Resources*

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Plant Latex, from Ecological Interests to Bioactive Chemical Resources* Published online: 2019-05-28 Reviews Plant Latex, from Ecological Interests to Bioactive Chemical Resources* Authors Luis Francisco Salomé Abarca 1, Peter G. L. Klinkhamer 1, Young Hae Choi 1, 2 Affiliations ABSTRACT 1 Institute of Biology, Leiden University, Leiden, Historically, latex-bearing plants have been regarded as im- The Netherlands portant medicinal resources in many countries due to their 2 College of Pharmacy, Kyung Hee University, Seoul, characteristic latex ingredients. They have also often been en- Republic of Korea dowed with a social or cultural significance in religious or cult rituals or for hunting. Initial chemical studies focused on the Key words protein or peptide content but recently the interest extended plant exudates, latex coagulation, mechanical defense, to smaller molecules. Latex has been found to contain a broad bioactive latex metabolites, endophytes, interaction range of specialized metabolites such as terpenoids, cardeno- lides, alkaloids, and phenolics, which are partly responsible for received February 28, 2019 their antibacterial, antifungal, anthelmintic, cytotoxic, and in- revised May 15, 2019 sect-repellent activities. The diversity in biology and chemis- accepted May 16, 2019 try of latexes is supposedly associated to their ecological roles Bibliography in interactions with exogenous factors. Latexes contain DOI https://doi.org/10.1055/a-0923-8215 unique compounds that are different to those found in their Published online May 28, 2019 | Planta Med 2019; 85: 856– bearing plants. Exploring the feasibility of plant latex as a 868 © Georg Thieme Verlag KG Stuttgart · New York | new type of bioactive chemical resource, this review paper ISSN 0032‑0943 covers the chemical characterization of plant latexes, extend- ing this to various other plant exudates. Also, the factors influ- Correspondence encing this chemical differentiation and the production, Dr. Young Hae Choi transportation, and chemistry of the latex exudates are de- Institute of Biology, Natural Products Laboratory, scribed, based on ecological and biochemical mechanisms. Leiden University We also proposed a latex coagulation model involving 4 gen- Sylviusweg 72, 2333 BE Leiden, The Netherlands eral conserved steps. Therefore, the inherent defensive origin Phone:+315274510,Fax:+315274511 of latexes is recognized as their most valuable character and [email protected] encourages one to pay attention to these materials as alterna- tive sources to discover metabolites with insecticidal or anti- microbial activity. This document was downloaded for personal use only. Unauthorized distribution is strictly prohibited. Introduction individual compounds but as a broad range of chemical classes. These chemical characteristics distinguish latex from other plant Latex is a milky fluid consisting of tiny droplets of organic matter exudates. In plants, latex is reported to be ubiquitous in more dispersed in an aqueous medium (a sort of natural colloidal sus- than 20000 species from over 40 families of angiosperms, which pension). Based on their physicochemical characteristics there represents between 8 and 10% of all of the terrestrial angio- are many subtypes of latexes, but the most well-known ones are sperms [3–6]. Among these, most studied plants belong to the rubbery latexes, of which polymeric microparticles can reach over Asclepiadaceae, Sapotaceae, Anacardiaceae, Apocynaceae, and 50% of the latex weight. If boiled, latexes can often coagulate, Euphorbiaceae families [1,7] forming a solid mass. The main constituents of the coagulum are The biological roles of latex are associated with many exoge- cis or trans-polyisoprenes and resinous materials [1,2]. Addition- nous factors. Because of their chemical selection and mutual in- ally, they also contain diverse primary and specialized metabolites as well as considerable levels of polypeptides and proteins. Com- pared with other types of plant exudates, latexes have a high de- * Dedicated to Professor Dr. Cosimo Pizza 70th birthday in recognition of gree of chemical diversity, not merely at the level of analogues of his outstanding contribution to natural product research. 856 Salomé Abarca LF et al. Plant Latex, from … Planta Med 2019; 85: 856–868 teraction of components related to their dedicated biological functions, plant latexes have been considered the first line of de- fense against many herbivores and pathogens [5,8,9]. It is possi- ble even that microorganisms living inside a host plant, so-called endophytes, could be involved in the production and biotransfor- mation of a broad range of host plant-originated metabolites re- lated to defensive strategies [10, 11]. Therefore, the complex fac- tors involved in the production of latex metabolites, together with plant biosynthesis, result in the chemical discrimination of the latex metabolome from that of other parts of the plant, showing a great potential as a new bioactive chemical resource. Looking into the feasibility of plant latex as a new type of re- source of active chemicals, this review paper deals with the chem- ical characterization of plant exudates in the first part and then later focuses on plant latexes and how their chemical composition generally differs from their bearing plants. Moreover, to investi- gate the factors influencing this chemical differentiation, the production, transportation, and chemistry of the exudation of latex are described from an ecological and biochemical mecha- nism perspective. Lastly, an insight for the use of latexes as a bio- inspirational model are discussed, which could partly explain me- tabolite selection, their complementation as bioactive mixtures in plants, and their potential application in medical and crop sci- ences. Characterization of Plant Latex ▶ Fig. 1 Plant exudatesʼ classification, their main chemical compo- nents and their producing structures. Pictures were taken by one of and Other Exudates the authors, Luis Francisco Salomé Abarca. In addition to latex, plants produce several types of exudates such as gums, mucilage, essential oil, oleoresins, resins, and phloem- like saps [1]. The differentiation of exudates is mainly associated families such as Fabaceae, Burseraceae, and Pinaceae are known with species specificity, anatomical origin, morphological appear- for their high resin production [1]. ance, or their chemical composition. Nonetheless, plant exudates Essential oils are mainly composed of organic volatile com- are generally classified based on their chemical composition pounds, which confer their characteristic properties such as odor (▶ Fig. 1). and density and low melting points [15]. Unlike other types of Gums are defined as solids usually produced from stem barks plant exudates, their vapor pressure under atmospheric condi- or roots. They flow out naturally upon mechanical damage or at tions is so high that they diffuse through the air [16,17]. These an infection point, readily solidifying on contact with air [2]. The volatile compounds belong to various chemical classes such as main constituents of gums are polysaccharides, which are alcohols, ethers or oxides, aldehydes, ketones, esters, amines, polymers of arabinose, galactose, mannose, and glucuronic acid amides, phenols, heterocycles, and mainly monoterpenes [16]. [1]. They are thus hydrophilic, being either water-soluble or capa- Extra fascicular phloem is a particular type of sap localized in a ble of absorbing water and swelling up to form a gel, but they are special network of sieve tubes outside of the vascular bundles. It is This document was downloaded for personal use only. Unauthorized distribution is strictly prohibited. insoluble in oils or organic solvents. found in a very limited number of families such as Curcurbitaceae Mucilages are water-soluble acidic or neutral polysaccharides [18,19]. They contain abundant amino acids and a wide range of that yield mixtures of simple sugars (e.g., galactose, arabinose, low levels of small molecules together with some proteins [20]. xylose, rhamnose, and galacturonic acid) or uronic acid [2,12, The biological functions of the phloem are likely associated with 13] by hydrolysis. The localization of mucilages is very specific, their proteins such as instant wound signaling and plugging of being produced intracellularly and having some of their constitu- the sieve elements to avoid the loss of nutrients. They have addi- ents related to cell wall components [14]. Differently to gums, tionally been found to play a direct role as anti-insect agents mucilages are strongly related to seed survival and also act as against generalist insect-herbivores [21]. water storage in plants [14]. The definition of latex has varied throughout time according to Oleoresins are referred to as resins or, more specifically, soft the review paper by Hunter (1994) [22]. Polhamus (1962) stated resins. They are exudates with a high content of volatile com- that “no one has demonstrated why a plant makes rubber and it pounds. The physical state of a resin varies from solid to semi-sol- does not appear to be a food reserve. Strong evidence indicates id, being composed of lipophilic terpenes such as mono-, sesqui-, that rubber is an end-product that is not reused in the metabolism and diterpenes. While most plants can produce resins, only a few of the plant” [23]. Five years later Metcalfe (1967) stated, “The Salomé Abarca LF et al. Plant Latex, from… Planta Med 2019; 85: 856–868 857 Reviews ▶ Fig. 2 A model of latex coagulation process. Four main steps are represented:
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