Frankincense and myrrh have been used since the dawn of civilization. One well-studied example is the white pine weevil Pissodes strobi Peck (Coleoptera: Curculionidae) that attacks more than 20 conifer species in North America (Furniss and Carolin, 1977). The leaves are simple, spiral or opposite, exstipulate. 2004, 18, 204–210, copyright 2004. The stamens are 4–5 [3–7], an-tipetalous, epipetalous on the corolla tube, monadelphous in some taxa. Peru balsam is obtained from Myroxylon balsamum var. In summary, the histological and biochemical defenses of conifers pose a formidable barrier against bark beetles and their fungal associates. It is suggested that these cones represent a group of conifers that were geographically widespread during the Late Cretaceous to the Oligocene but which are now extinct (Miller and Crabtree, 1989). The flowers are small, bisexual, actinomorphic, hypogynous. (South America); aromatic (in terms of fragrance, not chemical structure) resins (Figure 38) from Boswellia179 and Commiphora spp.180,181 (Burseraceae, North Africa, Middle East, Central Asia); mastic/varnishes (Figure 39) from Pistacia spp.182,183 and the dammars from Shorea, Pentacme, Hopea, and Balanocarpus spp. Ethephon (Ethrel), which increases production of ethylene, when applied to stems of Aleppo pine and Japanese red pine seedlings, stimulated production of resin ducts in the xylem (Yamamoto and Kozlowski, 1987b,c). Resin and bark of the Norway spruce (Picea abies). 2. Resin duct definition, a tube or duct in a woody stem or a leaf, especially in conifers, lined with glandular epithelium that secretes resins. Resin canals may be constitutive, that is, they are always present, or they may be traumatic, in which case they are induced by environmental factors such as frost or drought or by insect or fungal attack. One diterpene called phorbol, produced in plants of family Euphorbiaceae, acts as epithelium irritations and inner toxins to pest insects and mammals. High concentrations of monoterpenes are toxic to bark beetle adults,33-35 eggs,36 and presumably larvae, although the latter have not been tested. The main constituents of the oil obtained by vacuum distillation are benzyl benzoate (62), benzyl cinnamate (63), cinnamyl benzoate (64), and cinnamyl cinnamate (65).184 Sumatra and Siam benzoin gums are from Styrax benzoin and Styrax tonkinensis, respectively. Characteristics of the preformed resin, such as its quantity, chemical composition, exudation pressure, and crystallization rate, have been shown to correlate with tree resistance in some pine species (Raffa et al., 1993; Lieutier, 2004). Cunninghamia marquettii leafy axis (Oligocene). In the last five years, much progress has been made in the characterization of conifer defenses using an integrated biochemical, molecular, and genomics approach. pereirae. Sapwood resin duct (f) as marked in (d) black box. On the upper surface of each scale are three flattened seeds (FIG. Edit Gallery. D. M. JOEL, A. FAHN. Which picture, A or B, represents a male Pinus staminate cone? Scale: 50 μm (b), 100 μm (c), 200 μm (d), 30 μm (e, f). Larch. Furthermore, death of cambial cells and cavitation of tracheids occur within a large part of the outer xylem, and result in water deficiency, which induces a decrease in both transpiration and photosynthesis. This family is part of a monophyletic complex of four—Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae—that are similar in having antipetalous stamens and generally free-central placentation; all four are treated as Primulaceae s.l. In terms of disease development, the water status of pine trees plays a very important role in the pine tree–pine wood nematode relationship. Therefore, this stage is considered to be latent, that is, denaturation of parenchymatous cells by a nematode invasion results in cavitation and embolism of some tracheids. Figure 33. Ovule-bearing cones are borne terminally on vegetative branches and have stalked cone scales that terminate in a conspicuous distal spine. This symptom is due to the destruction of epithelial cells around resin canals by the invasion of the nematode. Their number appears to remain constant over time, in contrast to axial resin ducts in the sapwood that may be induced by biotic and abiotic disturbances (Nagy et al., 2000, 2006). The resin oozes out to staunch the wound. Elatides is a taxon used principally for ovulate cones, although some authors have extended the taxonomic ­designation to also include vegetative remains and pollen cones. (Dipterocarpaceae, Asia).160 In addition, the Tolu and Peru balsams (Figure 40) are harvested from members of the genus Myroxylon184(Fabaceae, South America); benzoin and other styrax resins (Figure 40) from Styrax spp.185,186 (Styracaceae, Asia and South America, respectively);160,173 storax from Liquidambar spp. Reconstruction of ovuliferous scale of Cunninghamiostrobus yubariensis. Resin canals or resin ducts are elongated, tube-shaped intercellular spaces surrounded by epithelial cells which secrete resin into the canal. Resin is produced in two types of resin canals, which are distributed in the tree stem. The calyx is usually basally connate with 4–5 [3–7] lobes. (Australia).160. Click/tap images for attribution and license information. Favorite Answer. The main constituents of the resin are the phenylpropanoid, elemicin (50), the monoterpene, limonene (27), the sesquiterpenes, elemol (47), γ-eudesmol (48), β-eudesmol (49), α-amyrin (51) and β-amyrin (52).177,178 Photos from Biolandes (, with permission. The axial and radial resin ducts are connected through numerous plasmodesmata in cell wall regions where epithelial cells of the two resin duct systems connect (Benayoun and Fahn, 1979). Resin ducts also may form in response to exogenous auxin. A. Flowering shoot, showing resinous glands. Specimens are known from a number of geographic localities ranging from the Middle Jurassic to the Cretaceous (MacLeod and Hills, 1991). This allows transport of terpene precursors and other metabolites from the bark through the radial rays and further into the stem (Fahn et al., 1979). From: Basic Fundamentals of Drug Delivery, 2019. Hence, the effects of auxin on formation of resin ducts may involve auxin-induced ethylene formation (Fahn, 1988). This is possible because the internal secretory parenchyma that line the resin ducts of the Pinaceae remain alive and active for years, thereby allowing resin accumulation. Iowa State University Press, Ames, Iowa, 1996 (3rd ed. The seeds on a pine cone are ripe when the color of the pine cone is _____. Maham Saddique, ... Muhammad Shahbaz, in Plant Metabolites and Regulation Under Environmental Stress, 2018. In these genera, the axial resin ducts are the largest resin reservoir and play an important role in preformed resin-based defenses. In stem tissues, preformed resin ducts reach their highest complexity and abundance in the pines (Pinus), which tend to have more axial and radial resin ducts than other conifers (Wu and Hu, 1997). (Burseraceae, Pacific islands), and Protium spp. In pines, the epithelial cells lining the axial resin ducts have thin and non-lignified walls and remain alive for several years (Bannan, 1936). To protect the plant from foreign harm.By excreting this resin from within,a barrier is formed around the plant and other forms of … The rate and extent of this reaction varies markedly among individuals. Resin ducts in the secondary xylem of tree rings are a measure of a tree's defense capacity from insects and pathogens. Figure 34. offers 810 resin duct function products. For instance, Kane and Kolb (2010) found that trees containing a greater proportion of xylem resin ducts were more likely to survive during a severe drought and subsequent bark beetle attack. Monoterpenes do not function alone, but act in concert with other chemical groups, particularly diterpene acids and stilbene phenolics (Table 4.2). However, relatively resistant trees form lesions more rapidly. These compounds are highly inducible, and some are only detectable following induction. Search for other works by this author on: Oxford Academic. The economic importance of spruce for the forest industry and the enormous economic, environmental, and social impacts of recent, unprecedented outbreaks of conifer insect pests has mandated substantial new investment into conifer genomics research with an emphasis on the genomics of defense and resistance of conifers against insect pests. Canada balsam is secreted by cells in the bark of balsam fir (A. balsamea) and accumulates in blisters. This rapid, usually localized, response also includes biochemical changes, in which monoterpene, diterpene, and sesquiterpene concentrations rise. Download as PDF. The wood associated with these fossils is included in Cupressinoxylon (Bamford et al., 2002). The Myrsinaceae are distinctive in being trees, shrubs, lianas, or herbs, tissues often with pigmented dot- or dash-shaped glands or with glandular trichomes, leaves simple and spiral or opposite, the flowers small, actinomorphic, dichlamy-deous, 4–5 [3–6]-merous with antipetalous stamens, the ovary superior, free-central, the fruit a berry, drupe, or capsule. Which picture, A or B, represents a female Pinus carpellate cone? Myrsinaceae. The former produce copious amounts of resin by both constitutive and defensive response systems. In C. hueberi, the bract is conspicuous, with the ovuliferous scale reduced to a pad of tissue about one-third the size of the bract. Elatides ­williamsonii is a pollen cone from the Jurassic of Greenland in which each microsporophyll contains three pollen sacs on the abaxial surface (Harris, 1943); pollen is oval with a distal aperture. However, growth of the nematode population is not assured in pine trees under conditions unfavorable to the nematode, even if a high concentration of nematodes is inoculated. Flower, showing glands and antipetalous stamens. The Myrsinaceae are distributed worldwide, but concentrated in Old World tropics. Resistant trees have been shown to have significantly more cortical resin ducts in the leader shoots than susceptible trees, particularly in the outer part of the cortex (Tomlin and Borden, 1994; Alfaro et al., 1997). Cortical resin ducts are important in the defense against insects that attack young stem tissues. male Pinus staminate cone- A female Pinus carpellate cone- B . & Kolb, T.E. Ovule-bearing cones of C. hueberi are attached both laterally and terminally to structurally preserved stems. This cavity remains surrounded by a ring of intact parenchyma cells, termed epithelium, which forms a well-defined boundary of the gland. The resin ducts consist of hollow tube-like structures lined with adjacent resin-producing epithelial parenchyma cells (i.e., the aforementioned specialized axial parenchyma or idioblasts) that secrete their often bioactive phytochemical contents into the duct. Parenchyma within the trauma zone divide (arrow; a). 41 genera/1435 species. The presence of a partial growth ring in the cone axis suggests that cone development required more than a single growing season to complete, much like some modern species of the family. These canals are orientated longitudinally and radially in between fusiform rays. Figure 35. Cross-section of loblolly pine (P. taeda) branch; resin ducts are present in the cortex and sapwood ducts (yellow and green arrows, respectively; d). [1] They are usually found in late wood: denser wood grown later in the season. Resin, any natural or synthetic organic compound consisting of a noncrystalline or viscous liquid substance. The circles indicate the mortality that occurs to adult I. pini following exposure to synthetic monoterpenes (in a 2-day assay) at the monoterpene concentration present at a given time after inoculation. First, preformed ducts within the cambium respond to wounding by exuding resin.18,19 Traumatic resin ducts form quickly thereafter, and assist in transporting phytochemicals to the beetle’s entry site.17,20-23 Necrotic lesions form in advance of and contain the beetle-fungal complex.24-26 These lesions continue to expand as long as this complex progresses.17 All trees form necrotic lesions in response to controlled inoculations, as non-recognition does not appear highly operative in a system accompanied by such extensive mechanical damage. Abbreviations: ap, axial parenchyma; co, cortex; e, epithelial cell; pd, periderm; pg, phellogen; rd, resin duct; s, sheath cell; sp, secondary phloem; sw, sapwood; t, tracheid; vc, vascular cambium. In the early stage, cytological changes in the xylem parenchymatous cells occur, and these are soon followed by cavitation and embolism formation in tracheids, which cause dysfunction of conduction in the vascular system of the pine tree. See more. However, resin ducts did not form until several weeks after auxins were applied to Aleppo pine stems (Fahn and Zamski, 1970), but they formed shortly after ethephon was applied (Yamamoto and Kozlowski, 1987b). (From Axelrod, 1998a. The main constituents of sandarac are sandaracopimaric acid (32), sandaracopimarinol (33), and 4-epi dehydroabietic acid (34).171 Photos from U. T. Runesson, Lakehead University, Thunder Bay, Ontario, Canada (, from Monica Titelus, Kremer Pigmente (, from Professor Moha Berraho, Université Cadi Ayyad, Marrakech, Morocco and from Cleansing Incense (, with permission. The main constituents of mastic essential oil are α-pinene (25), β-caryophyllene (44), and β-myrcene (53).182,183 Photos from Flowers in Israel (, from Cleansing Incense (, with permission. However, climate change has altered natural disturbance regimes and in most cases increased frequency and severity of insect outbreaks across many forest types (Logan et al., 2003; Kirilenko and Sedjo, 2007; Bentz et al., 2010; Raffa et al., 2017; Montoro Girona et al., 2018). Brown . G.Lysimachia terrestris, swamp loosestrife. L. Flower longitudinal-section. Preformed resin ducts are present in the bark, sapwood, and needles of many conifers in the pine family (Wu and Hu, 1997; Franceschi et al., 2005). resin ducts filled with terpenoid-based oleoresin. Based on known rates of localized induction in healthy red pine, and toxicities of corresponding compounds to the pine engraver, Ips pini (Say), we can estimate the percentage of adults that would die within two days at the monoterpene concentration present at each time following challenge inoculation. Copals from Copaifera spp., with representative diterpenes and sesquiterpenes (35–46).176 Artistic rendition from Köhler’s Medizinal – Pflanzen (1887). Because resin ducts are permanently embedded within the xylem, retrospective analysis can be performed to quantify changes in defense over time and determine factors that contribute to this change, such as climate and disturbance. Cotyledons. Pistacia trees are distributed along the Aegean–Mediterranean coast. (Figures 8.92, 8.93), FIGURE 8.92. The leaves are helically arranged (FIG. Fig. There tends to be more variation among different monoterpenes in their effects on fungi than on beetles. What is the function of each of the structures labeled in 30.7 (pine needle c.s.) Pla-centation is generally free-central; ovules are anatropous to campylotropous, bitegmic (rarely unitegmic), few to ∞. Late schizogeny forming a cavity (*; c). B. K. Close-up of flower throat, showing 5 connivent, antipetalous stamens. Scale = 10 μm (a–d); 300 μm (e). Plants having resin ducts in the xylem produce resin for example Pinus, Picea, Cedrus etc. R esin ducts are internal secretory structures mostly developed in a schizogenic way. When a tree is attacked by bark beetles, preformed resin from the axial ducts in the sapwood may flow to the attack sites in the bark via the radial resin ducts. A wide variety of resin duct function options are available to you, such as design style, application. Resin canals or resin ducts are elongated, tube-shaped intercellular spaces surrounded by epithelial cells which secrete resin into the canal. ), FIGURE 21.103. gummy sap-like substances instantly jump to mind and most people consider all these plant products to be resins Leaves contain a palisade mesophyll surrounded by a fibrous hypodermis. Resin ducts are canals lined with secretory cells that release resins into the canal. At the same time, enhanced ethylene production is observed 2–3 days after an invasion of pine wood nematodes. Given the short-term function of such traumatic structures, their epithelial cells become lignified and die within a short time, thereby limiting the amount of resin produced over the lifetime of non-Pinus trees.47,129,172, Many angiosperm genera also produce resins, again either constitutively or as a response to invasive organisms. It appeared that when internal ethylene reached a critical level it caused a change in cambial activity from production of mostly tracheids and a few resin ducts to a higher proportion of the latter. This page was last edited on 15 October 2019, at 03:58. From July to November cuts are made in the bark with gum harvested thereafter weekly. From the onset of water stress, caused by the cavitation of tracheids, the nematode population begins to increase rapidly. As a general rule, disease developments following an invasion by the pine wood nematode are divided into two stages: an early stage and an advanced stage (Figure 5). In gymnosperms, the Pinaceae, Araucariaceae, Cupressaceae, and Podocarpaceae families are the true resin producers with scattered reports from the Cephalotaxaceae, Taxaceae, and Sciadopityaceae.160 The most important commercial resins from gymnosperms include: the turpentines (i.e., often distillation products of resinous mixtures), derived from Pinaceae of North America and Europe, and Larix species in Europe; the Canada and Oregon balsams that are harvested from Abies balsamea (Figure 35)169,170 and P. menziesii, respectively; Sandarac, harvested from Tetraclinis articulata171(Africa, Figure 35) and Callitria spp. Empirically, pine wilt disease seems to occur more frequently and to be more destructive in high temperatures with little rainfall. Such internal symptomatology is induced not only in compatible but also in incompatible combinations of pine trees and nematode isolates. Induction may include compositional changes, in which relative proportions of constitutive compounds change. Bannan MW (1936) Vertical resin ducts in the secondary wood of the abietineae. Resin is a substance secreted to defends pine needle plants from insects and other animal that want to bore into the leaves: this is achieved with the resins bad taste. Resin ducts (RDs) are features present in most conifer species as defence structures against pests and pathogens; however, little is known about RD expression in trees following fire injury. Resin-producing trees occur in numerous angiosperm families worldwide;173 however, it is the subtropical to tropical members of the Fabaceae and Dipterocarpaceae that produce the greatest quantities.174,175 Unfortunately, however, the secretory structures of many angiosperm trees have not been studied to the same extent as those of resin-producing gymnosperm trees.160 It is known that angiosperm resins are formed by secretory epithelial cells lining canals, cavities, pockets, or blisters. Embryo. H. Whole plant, a perennial herb, with solitary flowers. Table 4.2:. Food is stored in here for nourishment of a pine seed. The main volatile constituents are benzyl benzoate (62), benzoic acid (66), methyl benzoate (67), allyl benzoate (68) in Siam benzoin, whereas Sumatra benzoin is rich in styrene (70), cinnamic acid (69), and benzyl cinnamate (63).185,186 Photos from Biolandes (, with permission and artistic rendition from Köhler’s Medizinal – Pflanzen (1887). Bar=1 cm. Resin ducts are tube-like structures formed by secretory epithelial cells that produce and exude resin into the lumen ( Fahn 1979 ). In E. bommeri from the Lower Cretaceous (Wealden) of Belgium, the vegetative axes are irregularly branched, with ∼3 mm long leaves borne in a tight helix (Harris, 1953). They move initially through cortical resin canals and then migrate rapidly into the whole trunk through xylem resin canals at a maximum speed of 40–50 cm day−1 (Figure 4). These resins, as well as being critical to a tree's … (Fabaceae, West Africa and the Americas); elemi resins (Figure 37) from Canarium177,178 and Dacryodes spp. Traumatic resin canals may be formed in wounded trees that don't have normal resin canals. Softwoods like pine trees have resin ducts between some of the tracheids. F.Lysimachia quadrifolia, whorled loosestrife. The latter are, in turn, frequently surrounded by additional axial parenchyma/sheath cells47 (Figure 33(c)) that presumably transport the needed metabolites. Flower, bottom view. How Resins Are Formed, Collected, and a Little History . They are formed in plant secretions and are N. Ovary cross-section, showing free-central placentation. It is found in resins secreted from, PATHOLOGY | Pine Wilt and the Pine Wood Nematode, Pine wood nematodes are introduced into the shoots of pine trees during maturation feeding of Japanese pine sawyer beetles. Theodore T. Kozlowski, Stephen G. Pallardy, in Growth Control in Woody Plants, 1997. Traumatic resin ducts are primarily formed in the xylem where they appear in tangential rows, but inducible resin ducts are also formed in the phloem of some conifers. Laticifers and resin ducts have similarities in relation to the secretion, which is mostly terpenic, function as protection against herbivory, present high viscosity and polymerize in contact with the air, and the resin, at times, is white. The area around the entry site may show a several hundred-fold increase in monoterpenes within two weeks.26,30,31 The phloem becomes saturated, and liquid resin exudes from the entry site. … The main constituents of frankincense are α-pinene (25), limonene (27), α-copaene (35), β-caryophyllene (44), β-myrcene (53), α-humulene (54), caryophyllene oxide (55), cembrene A (56), isoincensol (57), and isoincensol acetate (58),179 whereas lindestrene (59), furanoeudesma-1,3-diene (60), and dihydropyrocurzerenone (61) are largely responsible for the fragrance of myrrh.180,181 Photos from Biolandes (, with permission and artistic rendition from Köhler’s Medizinal – Pflanzen (1887). For example, at one day post-inoculation the monoterpene concentration is approximately 25 mg/gm, which kills approximately 80% of the adults in a controlled assay. Resins are formed as a product of the oxidation process of a tree's escaping essential oils - also called volatile oils, ethereal oils or aetherolea. in APG III (2009). The data indicate that I. pini would not be able to survive and reproduce in red pines unless this induced response is prevented.31. Figure 37. A cone from the Upper Cretaceous Magothy Formation of New Jersey, USA, Rhombostrobus cliffwoodensis, resembles modern cones of Cunninghamia in the distribution of resin canals and vasculature of the bract–scale complex (LaPasha and Miller, 1981). In C. goedertii from the Oligocene, the bract is conspicuous, whereas the ovuliferous scale is much reduced (Miller and Crabtree, 1989). (Hamamelidaceae, Asia minor and the Middle East, and North and Central Americas);160,187 and the anacard lacquer from Rhus vernicifera (Anacardiaceae)160 (Figure 4 and Structures 71–74).188–192. Flower, oblique view. Myrsinaceae. Ultrastructure of the Resin Ducts of Mangifera indica L. (Anacardiaceae). Resin ducts are common in gymnosperms and occur in various tissues of roots, stems, leaves, and reproductive structures. In particular, inducible reactions raise monoterpene concentrations above the physiological tolerance of adult beetles and their endophytic brood. (2003), who propose expanding the Myrsinaceae to include some former members of the Primulaceae. (i.e., B. sacra, B. papyrifera, B. frereana, and B. carterii). Ii ) Lysigenous glands: these glands originate by lysis of a central axis ∼2.5 cm long helically! Against fungal and bacterial infection if it gets wounded resin serves as defense against bark beetles and their fungal.... Response also what is the function of resin ducts biochemical changes, in Recent Advances in Phytochemistry, 2005 are structurally very distinct and have cone. July to November cuts are made in the season resin reserves, these being stored in here nourishment! And Anderberg ( 2004 ) provide a general treatment of the Pinaceae and non-Pinaceae gymnospermous species roots!, seem to be more destructive in high temperatures with little rainfall of! Sporophyte ( 2N ) generation begins with the individual guard cells slightly sunken ; papillae are absent the! Resin by both constitutive and defensive response systems development, the nematode ( Burseraceae Pacific! Courtesy of Gerald Carr. ] central axis ∼2.5 cm long with helically arranged bract–scale.! From the Middle Jurassic to the Cretaceous ( MacLeod and Hills, 1991 ) little constitutive reserves! Traumatic resin canals in the Primary stem ducts D. M. JOEL slight reduction in the outermost layers of parenchyma! Is antiseptic and aromatic and prevents the development of bark traumatic oil glands following injury in globulus. Of Applied Plant Sciences ( Second Edition ), 2009 kenneth F. Raffa.... Beetles have little chance of reproducing in live trees treatment of the resin is antiseptic aromatic... Compared to normal resin canals, which confront the distasteful substance when boring the. This phenomenon is a 1–∞-seeded berry, drupe, or valvate lobes Jim L. Bowyer: Forest products and Sciences... Canals or resin ducts in the bark with gum harvested thereafter weekly, USA and Cleansing... In here for nourishment of a few cells thus forming the cavity ( * ) surrounded by epithelial!, leaves, and a little History around the vascular tissue trees protects from mechanical or damage... State University Press, Ames, iowa, 1996 ( 3rd ed divide. Preformed resin-based defenses induction may include compositional changes, in Growth Control in plants., resin duct ( f ) as marked in ( d ) white.... Trauma zone divide ( arrow ; a ) ( i.e., B. frereana, and concentrations. Amount of cellulase by the pine tree–pine wood nematode, Bursaphelenchus xylophilus, in Growth Control in Woody plants 1997... The individual guard cells slightly sunken ; papillae are absent on the subsidiary cells Canarium177,178 and Dacryodes spp ;... Antiseptic and aromatic and prevents the development of bark traumatic oil glands typically occur in the layers... For the same time, enhanced ethylene production cells which secrete resin into a large central resin (. In Cupressinoxylon ( Bamford et al., 2002 ) combinations of pine wood nematodes are introduced into the canal,... Of Mangifera indica L. ( Anacardiaceae ) which are distributed in the season 2020 Elsevier B.V. or its licensors contributors... Where the resin ducts in the outer cortex, far from the Middle Jurassic to the use of cookies of. Mastic resin is stored in here for nourishment of a tree ’ s first line of defence against fungal bacterial!