SECRETORY STRUCTURESOF AROMATIC AND

MEDICINAL PLANTS

A REVIEW ANDAnAS Of MICROGRAPHS

Authors: Katerina P. Svobodaand Tomas G. Svoboda

Micrographs: Andrew D. Syred,Microscopix photolibrary

Designed and Edited: Polly M. Syred

1\('''' to imu!:t',\ on front cover:

Top to bottom: Worm wood.Peppermint. Marjoram . Rosemary.

Gre ek O re-gano. LavenderBackground : Euca lyptus lea f

MICROSCOPIX PUBLICATIONSMiddle Travelly, Beguildy, Knighton . Powys t07 lUW. V.K.

Tilt' pffhlitllen 111111..' 110 representation, expressed fir implied. with regurd to theac( 'w'(/( y o/lhe ill!iwlI1alitlll contained ill IIIi,1 hook 111/(1 e(//IIWI O

PREFACE

Aromatic and medicinal plants arc used in the food . flavour, perfumery. pharmaceutical andhealthcarc industries. The ir usc is expected to increase every year due to a consumer-led trendtowards natura l foods. food and cosmetic additives and complimentary med icine includingaromathcrapy. Many branches of industry are searching for alternative. cleaner antimicrobial,antibiot ic. antioxidant and crop-protection agents. The possibility of utilising volatile oils isbeing investigated as it has been known since antiquity that they possess biological activities.During the last decade many books have been written about herbs and spices. chemicalconstituents found in various plant families and the ir uses.

However, very few research articles deal with the description of the sites where these chemi calsare produced. either on the surface of the plants or internally. There is a distinct lack ofillustrations whic h wou ld help the reader to visualise these anatomical structures. An earlyacquaintance with micrographs of secretory tissues increases the appreciation of the rela tionshipbetween the plant and its biochemical products.

This book is a source of usefu l tenus and concepts for descriptio n of a variety of secretorystructures. It does not cover the subject exhaustively but presents numerou s examples andexamines a few in detail. The reader is made aware of the end less variability in anatomical fonnand will therefore be better prepared to interpret the structure of other plant species and relatethem to those described in this book. An exhaustive list of references collected over the last tenyears gives the reader guidance for scientific approach to this area. Use of increasingly moreadva nced methods ofmicroscopic studies has resulted in remarkable insight into plant anatomy.diffe rentiation and development.

All photomicrograph s are original. prepared by Microscopi x from authentic plants and material.My botanical and phytoch emical interest. direct ed towards research on aromatic plants.unavoidably influenced the presentation of individual plant species. The problems ofclassification are not resolved and the order ofplant presentation is not related to taxonomy. Theplants arc presented in alphabe tical sequence of their common names, supporting interpretationsand conclusions that have been placed in the text. This method of organisat ion is simple, coherentand quickly per mitsthe reader acce ss to speci fic plates.

This book aims to be useful to both students and sc ientists with an interest in aroma tic andmedicinal plants and may be a valuable resource to all those involved in the research,development. cultivation. marketin g and uses ofthese crops.

Dr. K. P. SvobodaMev 2000

'Apricots are ripeWith moonli ght dripping

On each fruit'

Haiku by Shibata Hakuyojo

3

PLATE I

Mobile distillation unit being UM-'J for chamo mile oil extraction (Norfolk. Anglian Essential Oils Ltd.]

CONTENTSra~e no:

Preface 3l ist of plates _ 6Introduction _ 7Secretory structures in plants 9

Basil.. 15Bog myrtl e 16Caraway _ 17Catmint 18Cedar 19Chamomile. Roman 20Citrus 21Clove 22Cumin 23Dragonhead 24Eucalyptus 26Frankincense 27Ginger 30lIyssop 3 1Juni per 32Korean mint 33l avender 34l emon balm 35l ovage 36Marjoram 37Myrrh 28Nutmeg 38Orchid. lady's slipper 39Oregano 40Oregano. Greek 4 1Peppermint 42Perilla .43Plantain 44Rosemary 46Sage .47Tansy .48Wormwood 50

Bibliography 51List of plant species used in a ruma therapy 57Glossa ry 58Index of spec ies 60

,

Plate no:

2345&67 & x9& 10II & 1213. 14 & 1516.1 7.IX&1920.2 1 & 2223, 24 . 25 & 2627, 2R. 29 & 303 1 & 323334 & 3536. 37. 3S. 39 & 4041...2 3 & ......... 5 6 & 4748.49 & 5051 & 5253 & 5455. 56 & 5758.59 & 6061 &6263 &6465.66&6768 & 6970,7 [ & 7273 & 7475 & 7677. 7X. 79 & so8 182 & SJ84.85 & 8687 & ss89

90 .91 & 929394

LIST OF PLATES

P..~e no:Mobile distillation unit being used for chamomile oil extraction(Norfolk. Anglian Essential Oi ls ltd. ) .....Collection of various mint spec ies. ex peri mental plo ts at Auchincruivc XEcninacae angustitotia, experimental plots at Auc hinc ruivc 13Takeda . Japanese pharmace utical company herb garde n. Osaka 14Basil 15Bog myrtle 16Caraway 17Carmir u 18Cedar 19Chamomile 20Ci trus 2 1Clove 22Cumin 23Dragonhead 2...Detail ofdragonhead tlowc showing distribution ofsecretory glands 25Eucalyptus 26Frankincense and Myrrh 274 28A young Som alian woman grading (1-6) freshl y-collected resin for traders 29Ginger 30Hyssop 3 1Juniper 32Korean mint 33l avender 34Lemon balm .35Lovage 36Marjoram 37Nutmeg 3XOrchid. lady's slipper 39Oregano .40Oregano. Greek .4 IPeppermi nt .42Perilla .43Plan tain 44AgastacheJi.)('"iClllllm at Auchincruive herb garden 5Rosemary 6Sage 47Tansy .41\Experime ntal p lot of tan sy grown for volatile oil wi th high thujon c conte nt(A uchincruivc ) .49\Von11\\'oOO 50Street ma rket with herbs and spices . Seoul. Korea 56Sta tue of a stree t sell er w ith various herbs. essential oils and perfumes(Perfumery Museum. Grasse. South of France} 61

INTRODUCTION

Plants and their extracts have been used for centuries to relieve pain. aid healing and killbacteria and insects. They have also been employed in perfumery. cosmetics and relig iousrites. Plant chemistry became an es tablished discipline in universities at the end of thenineteenth century and since then many new structures have been discovered . The numberofnatural products obtained from plants has now reached over 100 000 and every year newchemical com pounds arc being discovered. Although their functions in plants have notbeen fully esta blished. it is known that some substances have growth regulatory propertieswhilst others are involved in pollination and seed dispersal. Many arc important asantifungal and antihcrbivorc agents with further compounds being involved in thesymbiotic associations in plants. Detailed information on indiv idual plant chemicals isava ilable in the Dictionary ofNatural Products and the Phytochemical Dictionary:

The complex nature of these chemicals, which arc usually prod uced in various type s ofsecretory structures . is also influenced and controlled by genetic and eco logical factors andsignificant ly, by the mode ofextract ion from the plants. The type of secretory structure is animportant charac teristic of a plant family. Detai led anatomical descrip tion of thesestructures is relevant to the market value of the plants. the verificat ion of authenticity of agiven species and for the detect ionofsubstitution or adulteration. It also provides a guide tothe method ofprocessing.

Microscopical investigation of plant structu res is an important part of the co mplexbiological research process which includes plant growth and developme nt. genetics andbreedi ng. There is endless variability in fonn and struc ture; observing these with themicroscope allows us to discover Nature in one ofits most powerful form s.

Plant material received forthese investigations was examined fresh or as chemically-fixedand stained sections for light microscopy (LM), chemically-fixed and cr itical point dried(CPD) for scanning electron microscopy (SEM) or cryo genica lly fixed for cryo-SE M. Nosingle technique was ideal for all purposes since each had its ovvn limitat ion and risk ofintroducing art ifacts but by employing several methods these difficultie s were reduced togive a clearer understanding of the structures. For LM. thin transverse sections (TIS) werecu t from wax-embedded material and stained to provide colo ur contrast. however, theinvolved chemical processes invariably caused some damage to the samp le. This wasalleviated by examining fresh. unsta ined sections either by differential interferencecontrast (Ole) or, ifsufficient contrast was atta inable. bright field illumination (BF) whereno chemical or staining processes were employed. SEM allowed high magnifica tionstogether with improved depth offield giv ing an overall view ofthe struc tures. Since SEM isincapable of recording colour. all images produced using this meth od were manuallyenhanced using computer techniques.

7

PLATE 2

. > -rimcntal plots al AuchincruiveCollection of'various mmt ...pecres. 1.:\P'

,

SECRETORY ST RUCTURES II\' PLA NTS

Plant chemica ls can beclassed as primary or secondary metabo lites. dependi ng on whet her o r notthey have an essential role in plan t metabolism and arc un iversall y present in all plants. Primarymetabolites include sugars. amino acids. nucleic acids and the chlorophylis. Secondarymetabolites mak e up all the remaining plan t chemicals from alkaloids to phen ols.

Essential oil s and other secondary plan t metaboli tes arc fou nd in a wide range of pla nt speciesincluding annual, bienni al or perenni al herbaceou s plants. eve rgreen or deciduous shrubs andtrees. The eco logica l and evolutionary role of these secondary me tabolites has been assoc iatedwith defence again st anima ls. healin g of plant orga n wou nds. protect ion from harm ful insec ts.resistance to microbi al attacks and attraction of insects and anima ls fo r pollinat ion. Severalspecies and varieties of plants. mostly those of commercia l interest. were investigatedsys tematically and in dep th. Recen tly. vario us stud ies co ncerned with secretory struc tures andfactors influencing their developmen t have been undertaken by research groups in biologica l andpharmacological departments.

Secretion is a commo n feat ure of living ce lls and invo lves the discharge of substances to theexterior (exotropic secretion) or into special intercellu lar cav ities (endotropic secretion). Thesearc spec ialised cells and the secreted material may co nta in various salts. latex. waxes. fats.flavonoids, sugars. gums. mucilages. essential oi ls and resins. It has been assum ed that theseprod ucts arc biosynthesised insitu and direct evidence for the biosyn thct ic ca pac ity ofg land ce llshas become available relatively recent ly with the development ofprocedures for gla nd isolation.These method s have yielded defin itive proof of the presence of enzymes speci fica lly withingland ce lls.

Trichomes present on the leaf surface and other secreto ry tissues can be examined using light.scanning and transmi ssion elec tron microscopy wh ich enables detailed ob servation of majorstages in the development of secretory cells . including their membrane system and nuclei . theoverall s ize ofthe gla nd and the amount of matcrial released into the subcuticular cav ity.

Essential oi ls. with or without accompanying resins and gu ms. are most commonly found inspecial sec retory structures either on the surface of the plant or with in the plant tissues. The typeofstructure is family or species speci fic. This ca n be usefu l in ident ificati on of plant material andveri fying the authenticity ofthe plan t source in the case of suspected adu Iteration.

St:CH.ETOR,\CI::U.S

The most simple secretory struc ture is a single secretion-containing ce ll where it is only the actualcontent that distin gui shes it from adjacent non-secretory ce lls. However, it may also be largerthan the other cell s or have a thick cuticularized lin ing. This ce ll type is found in many differen tplant tissues: in the leaf parenchyma of lemongra ss tAndropogon spp.) . bay (LaUrElS nohilis) .citronella (Cymbopogon nardus and C. winterianusv and patchouli (Pogostemon patchouli). inthe seed coat o f cardamom (E lellaria cardamonunn) , in the rhizome of g inger (Ziflgiber

9

oj]icinale) [Plates 42-44] and turmeric (Curcuma longa), in the fruit wall of pepper (Pipernignlm), capsicum and chillies (Capsicum annllm), in the pcrisperm and embryo of nutmeg(Myristica fragrans) [Plates 63 & 64 ], in the bark of cassia (Cassia angustifolia} and cinnamon(Cinnamomum zeylanicum}and in the root of valen an (Valeriana officinaliss,

O S.\IOPII OR [ S

Osmop hores are areas of flower tissues with secretory cells differing structurally from theadjacent cells (e.g. isodiametrie cells in orchids).

S IT RI-:TO RY C \ VIT IES

These cavities are more or less spherical structures that can be formed in two ways: theparenchyma cells can separate one from another leaving intercellular spaces called lumina orlacuna, or an actual cell can disintegrate leaving a cavity within the tissue. These spaces are linedwith secretory cells or an epithelium that produces the essential oils. In high oil yielding plantsseveral layers of these secretory cells are fonned. The cavities continually enlarge and somebecome filled with cells with thin, convoluted walls which also store the oil produced from withintheir plastids . Included in this group are fruits and leaves of plants in the Citrus family (c.ourantifolia. C. aurantium. C. bergam ia, C. sinensis. C. limom [Plates 20-22] as well asEucalyptus spp. [Plates 34 & 35] and buchu leaves. Citrus peel oils arc confined in oblate tospherical-shaped oil cavities (glands), sometimes cal led oils sacs, that arc located irregularly in theexoca rp of the fruit [Plate 22]. These cavit ies have no walls and are embedded at different depths inthe flavedo (the coloured outer portion or skin of the fruit). The glands of grapefruit lie deeply inthe flavedo and those of mandarin are likely to be nearer the surface. Fruits and leaves of theseplants are covered by a thick cuticle which is waterproof and also the primary means of waterconservation. Being shiny and reflect ive it is capable of deflecting some of the excess solarradiation in tropical and subtropical regions; it also reflects ultraviolet light. thereby protect ing theDJ A from the mutagenic effects ofsunlight. It is an excellent protect ion against fungi and bacteriasince they have no enzymes capable of digesting cutin . Secretory cavit ies arc also present in theflower buds ofcloves (Syzygium aromaticum) [Plates 23-26] , the fruit walls ofpimcnto (Pimentodioica), and in the elongated cavities in the bark ofmyrrh (Commiplwra molmo f), benzoin (Styraxbenzoim and frankincense (Boswelttaspp.) [Plates 36 & 37].

SEC RETORY Dl J(.. S

Ducts are elongated cavities. They can often branch to create a network extending from the rootsthrough the stem to the leaves, flowers and fruits . They are composed of an epithelium whichsurrounds a central cav ity. Several predisposed cells within the parenchyma undergoasync hronous division and in doing so they expand the initial space in the middle where the cellsarc all adjacent to form a cavity. Some ofthese cells formi ng the wall of the cav ity willchange intosecretory epithelial cells. The oils are biosynthesised within their leucoplasts and move via theendoplasm ic reticulum into the cavity. These cavit ies then becomejoined to form duct s. They canbe found in all of the Umbe lliferae family including anise tPimpinella anisum}; fenneltFoeniculum vulgares. dill tAnethum graveolenss. coriander (Coriandrum sativunu, cumin

10

(C"minum cymillum) [Plate s 27-30]. parsley tPetmselinum crispum) and angelica (Angelicaarchungelica s. In the case of celery (Api um graveolens s they can branch to create a networkextending from the roots. through the stem to the leaves. then to the flowers and finally to thefruits where they are known as virtac (sing. vitta). They are also present in the Pinaceae 1Plates13- 15], Compc sitac. Hypcricaceae and Coniferae families. The resin ducts in the xylem ofConiferae can reach 4-10 em in length with between 2 and 7 ducts per leaf.

G1..-\:\ I>lTL-\ K TK IC IIO \ IES

Glandular trichomes arc modified epide rmal hairs and can be found covering leaves, stems. andeven parts of flowers such as the calyx in many plant s of the Labiatae family. These include basil(0('imu11I basilic/lm) [Plates 5 & 6l. lavender (Lavandula spp.) [Plate 53 & 54], marjoram [Plates61 & 62] and orega no {Origanum spp.) [Plates 68-72J, mint (.I1elllha spp.) [Plates 73 & 74] andthyme (Thymus sp.). The secretory cells arc attached by a single stem or basal cell in theepidermis. The outer surface of the gland is heavily cutini zed. A toughened cuticle in which nopores or perforations are present. usually com pletely covers the trichome [Plate 7). The essent ialoils accumulate in subcuticular spaces and it is thought that they diffuse outwards through thecuticle. The glandular cells di ffer from normal plant cells in that they have a very large nucleusand a dense protoplasm that lacks a large central vacuole. There are numerous plasmodesmataacross the walls of the gland cells especially between the stalk cell and the collecting cell. In thevery young gland the intracellular organisation is almost identical to that of the adjacent cells butas the secretory cells develop complex changes occ ur. The membrane system progressivelydegenerates and in thc fully-developed glands only a fine granular cytoplasm remains.

Cells in the multicellular heads usually have nucle i with double the norma l numbers ofchromosomes (endopolyploidy) and in fully-mature glands. mitochondria are the most abundantorganelles which migh t reflect a high energy requirement. There is also a well-developedendoplasmic ret iculum. Essential oils accumulate in the subcuticular cavity [Plates 32, 70 & 92].All these changes in the glands occur at a very early stage of the leaf development and theglandular hairs are fully-fonn ed by the time the leaves are about 5 mm long. The formation andtransformation of the essential oils. however, takes place continuously from the formation of thegland unti l senescence. The essential oil biogenet ic precursor isopentenyl pyrophosphate is. with'iigh probability, synthesised in the leucoplasts.

Therearc two main types ofglands that canexh ibit minor var iations:

(i) Pcltate glands with one basal cell. a short stalk and a large six- to eight- celled head[Plates 61, 84 & 86J.

(ii) Capitate trichomes with either -

a) one basal cell. a short monocellular stalk and a two-cellular head [Plates 72 & 85J.or

b) one basal cell. a mult icellular stalk and a small globose. unicellu lar head [Plate 67J.

II

In every species there arc distinctive variations in the size and shape of the glands. For example ,yarrow (Achillea millefoliumv secretory trichomes of the floret produce azulene and occurmain ly in the coroll a lobes of the ray and disk florets and also on the leaves. When a floretreaches 0.250101, fully-mature trichomes are present having developed from protodermal cells .They have 10 cells includ ing a pair of basal cells. a pair of stalk cells and 3 pairs ofglandularcells. In Cannabis sa tiva a variety of glandular trichomes occur, including bulbous (smallglands with a single-ce lled head and a unicellular stalk). capitate-sessile (a gland with a head of8 or more cells on a very short stalk) and cap itate-stalked (similar to previou s type but with asubstantial multicellular stalk). These glands arc considered to be the main location of thenarcotic cannabinoids.

It is poss ible to isolate an individual gland from the leaf surface and analyse its content forchcrnicalcomposition and presence ofspecific enzymes.

1':1'1HElnt" I. C[LLS

Essentia l oils obtained from flowers arc not usually secreted by glandular hairs but merelydiffuse through the cytoplasm. the cell walls and the cut icle to the outside. The yie ldofesse ntialoils from these species is generally very low. Examples include rose (Rosa spp.), 0.075% (w/v).acacia (Acacia spp.) 0.084% (w/v) andjasmine (Jasminum spp.)0.04% (w/v) .

Buds of a number of plant species (e.g. Aescuius. Alnus. Betula. Populus. Prunus andRhamnus ) also secrete lipophilic substances. mainl y flavonoid aglycones mixed with essentialoils. Secret ion here occ urs from epidermal cell s which are covered by a cuticle. The secretedmaterial is first eliminated into a space formed between the outer walls of the cells and thecuticle covering them, forming a blister that subsequently bursts.

C O:'\C LUS IO.\

Plants have prov ided man for hundreds ofyears with many of the basic and important materialsrequired for day-to-day living. including oxygen. food. clothing and timber. as well as being asource ofcompounds such as oils, resins, rubbers . gums. dyes. pesticides and drugs. Plants canbe considered as biochem ical factories that have been evolving their programmes over the last400 million years. Researchers have been elucidating chemical pathways and variousindustrial applications of these products, and anatomical desc ription is one of the mostimportant aspects ofeconomical uses ofthese plant s. It is also important to highlight the need tosafeguard and conserve the diversity of our global flora and to contribute with research intocareful exploitation of plant resources.

PLATE 3

Ecmnacae angustifolia , experimental plots 31 Auchincruive

13

PLATE 4

Takeda, Japanese pharmaceutical company herb garden. Osaka

BASIL - Ocimum basilicum

Plate s:Sessile secretory gland on 10\\cr leaf surface covered by turgid. intact cuticle (SE~f.CPD)['(559].

Plat e b: Leaf (T!S) showing sess ile secretory gland with colla psed. ruptu red cuticle. Chloroplasts visible inthe underlying. parenchyma (U,f. BF. fresh. unstainedt lx575].

15

-BOG MYRTLE - Myrica gale

Plat e 7: Sessile sl,..crcrory gland on 10\\cr leafsurface covcrcd by turgid. intac t cuticle (S E~t.CrO) [x60 7].

Plat e 8: Stalked glands and non-secretory trichomes on lower leafs urface (SE~l .CrD ) IxhOOj.

CARAWAY - Carum carvi

Plat e 10: Seed pair(T/Sl showing.f primaryand2 commissural \" ittac(SEr-. t, CPO) (x.fXJ.

17

CATMIl"T - Nepeta cataria var. citriodora

Plate II : Sess ile secretory g land on lower leaf surface covered by turgid. intact cut ic le. Part of non-sec retoryhairvisihlc in fo rcgroun d tSblvl. CrD)[x94~J .

Pla te 12: Sessile secretory gland on lower leaf surface with ruptured cuticle revealing underlying secretorycells (SE~LCPO) [x 1.250].

Platt' IJ : Needle (T iS) of C.atlantica with detail of resinduct (rd). Duct is lined withepithelium of secretory cells(5C) (LM, BF. stained withsafranin and fast green) [x333].

Plat e 14: Needle (T iS) of e.brevifolia with detail of anarrow resin duct (rd}(U.1, I3 F,stained with safranin and fastgreen) [d25].

Plat e 15: Wax-coated needle(T iS) ofe. atlantica showing 2well-formed resin ducts {rd].parenchyma and centra lly-located vascular tissue (vt).xylem and phloem ( L~l, BEstained with snfranin and lastgreen) [x64].

CEDAR - Cedrus spp.

CIIAl\IOl\I1LE, Roman - Chamaemelum nobile

Plate 16: Individual floret from composite flower con-red w ithsess ile secretory glands (S Hot. CPO) [xX3].

Plate 18: Upper leaf surface with sess ile secretory glands and 000-secretorytrichomes(SEM. CPD)[x1441.

Pla te 17: Detail of sess ilesecretory gland on floret surfacewith oil-filled subcuticular spacere su lt ing in fu lly-ex te ndedcuticle. Secretory cells visible atbase of gland struct ure (l M. BF.fresh. unstai ned ) [x297).

:'\ole: Cha mazulene. produced inthe oi l during distillation byth e rma l deco mpo sit ion ofmatricin. is responsible for theblue colourauon of the extractedoil - in its raw state the oil isvirtually co lourless .

Pluto 19: Developing ovule offully-mat ured flo wer coveredwith sess ile secretory glands(SEM. CPD)[x72].

C ITRUS - Citrus spp.

1)lalc 211 : Detail of secretorycav ity (TIS) in the flavedo oforange fruit JX'd (SEM. CP D)[x96].

Plat t 21: SCCrCIOl) ' cavit ies ,------------------------~(TIS) in the flavedo of orangefruit~I (LM. BF. stained withsafranin and fast green) (:

CLOVE - Syzygium aromaticum

Pla te 25: Dried stem (TiS) showingthe distribut ion of endogenous oilglands beneath the surface ( S E ~1.CPD)[xJ7].

Plate 26: Detail of endogenous oilglands (TIS) in dried stern (SEM,CPDI(x I39].

Plat e B : Dried petal (TIS) sho.... ingprolific occu rrence of endogenous oilglands (SE~ I.CPO) [x22~ J.

Plate 2~ : Detail of an endogenous oilgland (TIS) in dried petal (5E\ I, CPO)(x338) .

CU!\lI N - Cuminum cyminum

Plate 27: Detailof SCI,.'d pair (TIS) withcarpophcrcs and commissural vittac(SEM. CPDI ['91).

Pla te 2M: Seed pa ir(T/S)(SE~I . CPO)Ix:!J ).

I'l ale 29; Seed (TIS) showing 4primary and 2 commissural vitracsituated beneath the seed coat (S E~1.CPD) (xJt.)J.

Plate JII: Seed (TIS) showing. detail ofvittae (SEM. CP()[x147).

23

DRA GO:-lHEA D - Dracocephalum moldavica

Plat e 3 1: Scs"i lc srclory gland on lower l~afsurfacc ; stomata "sible on cpidcnnis (SEt\1.CPO) [x790).

Plate 32 : Sessile secretory gland on calyx (TiS) with oil-filled subcuticular space resulting in fully-extendedcuticle. Secretory ce lls visible al base ofgland structure. Non-secretory trichomcs present on right hand sideIL\ 1. RF. fn..sh, unstained) [:

PLATE 33

Detai l ofdragonhead 110wer show ing distribution of secretory glands [x::!O]

25

EUCALYPTUS - Eucalyptus citriodora

Plate Jof : Lcaf(T/S) showing secretory cavities. Oil removed by chemicaltreatment during preparat ion oftissue for microscopy rStixt. CPD)[x236].

Plat e .' 5: l caf (T/S)sbo ..... ing secretory cav il)' IL~1 . BF. sta ined with satra nin and fast green}[x23 1].

FRANKINCENSE- Boswellia carteri

Platt' _' it: Five year old stem (T iSI . Xylem and bark tissues clea rly visible with resin globules oozing fromducts t cryo-Slix! ) [xI18].

Prat e 37: Fi\ c year- old stem (T/S ). Xylem and bark tissue clearl y visible showing resin due ls in bark (fresh. unstain ed) [x-rr ].

FRA:\' K INCE:\'SE- Boswelia carteri

Plait' 38: Resin sample. maydigrade 3-4. sorted by localSomal ian collectors.

P131t' 39: Res in sam ple. maydigrade 1-2. sorted by localSomalian collectors.

Pla te ~O : Freshly-collectedmyrrh (Commiphnra myrrha)resin.

"

PLATE 41

A young Soma lian .... oman grading ( 1-6) freshly-collec ted resin for traders

GINGER - Zingiber officinale

Plate ~ 2: Rhizom e ( T IS)~how inlJ oi l globule ooz mgfrom secretory cell. Starchgranules also d early visible(cryo-S EM and etched) [x600] .

"lut e ..& 3: Rhizome (T I S)s how i ng oil g lob ules insecretory cel ls of parenchymatissue. Starc h granules abundant(LM, HF. fres h, unstained)[, 205J.

Pfat e ...... : Rhi zo me ( T I S)showi ng oil gfobules withinmembrane of secretory cell(cryo-S E ~ I and etched) I'(8 13).

30

HYSSOP - Hyssopus officinalis

I' late 45: Leaf (TIS) showingsessi le secretory gland on uppersur face wit h o i l - filledsubcuticular space resuhing infully-ex tended cuticle (LM. BF.stained with safranin and fastgrccntjx-r l l J.

Plate 46: Sessi le secretorygland with ruptured cuticle onlower leaf surface . Freshmaterial was dried at 80"C todemonstrate the destructiveeffect of hig h d r y i n gte mpe ratu res (SE M. C PD)[x 1.069].

(

Plat e -l7: Leaf (TIS) showingsessi le secretory gland on uppers u r face w it h o il - filledsubcuticular space resulting infully-extend ed cu tid e (LM. I~~l.,;:-':''';:~~!;~:'''",,:: ~':::':''':_'';:~.JOle. fresh. unsta ined )( x400].

JI

JUN) PER - Juniperus C0l1 l111Wl is

I' la ll UI: Nccdlc (T/S )showmg ccnuu l resinduct (rd)(Lt\.1. I3 F. stained with safmnmand fast grc~n )l x67 1 .

I' tate ~9: Gree n 1:11.'ny (TIS) showing sc'(r~lOry cavities(D.l. OF. sta ined with safra nin and fast gr~~n)lx~41.

Plate 5U: Ripe ben)' (TiS) detai l of ep ithelium-lin...-dsc-crctory cavity IL~1. RF. stained with safrunin and fast

grc~n ) lx~OOl

KOREAN MINT - Agastache rugosa

Plat e 5 I : Sessile secretory gland on lower leaf surface: stomata visible on epidermis ( SE~ I .C PO I[x 1,325J.

Plait' 52: Detail o f sessi le scc retory gland on lower lear surface (TiS) with single basa l cell. ~ ....-crctory cellsand oil-filled. partially-extended subcuticular space (D .I. BF.stained with fast green) Ix6(0) .

H

LAVEN DE R - Lavandula angustifolia

Pla it' ~3: Detail oflower leafsurface showing sessile secretory gland and non-secretory trichomes associatedwith water-conservation (SEI\1. CPO)[x638].

l'Iat e 54: Lower leaf surface show ing two types of sessile secretory glands and dense cove ring of non-secretory trichomes(SEM.CPO)(x251].

LEMON BALM - Melissa ojJicinalis

Plair- 55: l eaf (TIS) showingsess il.: secretory g land on lowersu r fac e (Uvl , Bf', fres h,unstai ned) [:d75).

Plat e 56: Genera l view oflowe rleaf surface showing sparsedistribution of sess ile secretoryglands. Extremely [ow densi tyc f trichomcs 10 per mm') isresponsible for low yield ofvolat ile oi l in this species (l M.incident illumination) [x4 1].

I'lale 57: Young leaf (4 weeksold) (TIS) with well-develo pedsessile sec retory gland filledwith oil. Non-secre tory hairalso present on upper surface( 1.1,,1.DIe ) [x370].

35

LOVAGE - Levisticum officinale

rl aft"\ 58 ~'l 59: Leaf at midrib (TIS) showine two oil ducts lined with secretory cells. Xylem. phloem andcambi um form the midrib vascula r tissue (L\l. SF, stained with safranin and fast green by tw o differenttech niq ucs jod- o il d uel; ph - phloe m: sc - secretory ce lls; xy - xylem [x-ltoJ; x4 10].

Plute 60 : Overall view orlcaf at midri b (TIS) with two oi l ducts c learly visible ILJ\t. BF. stained with safraninand fast green)(:

MARJORAM - Origanum marj orana

Plat e 6t : Sessile secretory gland on lower leaf surface (TIS). Four secretory cells clearly visible beneathruptured cuticle (LM. BF. stained with safranin and fast green)[x I.XX?].

Pla te 62: Stalked gland with unicellular head (TIS)and turgid. intact curictc on lower leafs urfac e ILM. Die.fresh. unstained) [x1.800].

37

NUTMEG - Myris tica fragrans

Plat e 63: Embryo (TIS) showing inner pcrispcrm and secretory cells. Preparation for CPO process hasremoved allo il from cell" (5E\ 1.CPD) Ix79}.

I'Jall'~: Detailof secretory cells (TIS) in embryo (SEM, epD) [x3S0}.

ORCHID, Lady's slipper - Cypripedium pubescens var. calceolus

Pia te 65: Stalk ed glandu lar trichomes 0 11 upper leaf surface (Uv1. incident illumination, fresh. unsta ined)[x I00].

Plate 66: Root (TIS) showi ng cortex parenchym acel ls ti lled with stnrch grains fSb lvl.C PD) [x67].

Platt.' 67: Leaf (TIS) showing stalked glandulartrichome with unicellu lar head (LM, BF, stainedwith safranin and fast green) [x205 ].

OREGANO - Origanum vulgare

Plato 6H: Sessile secretory gland on lower leaf surface. Impression of cell sutures nol visible on cuticlesurface due 10 distention byoil secretions (S H I. CPO )[x 1.1-'5] .

Pla te 69: Sess ile secretory gland on lower leaf surface with ruptured cuticle revealin g mdiv idnal secretorycelts. Neighbouring stomata clearly visible ( SE~I . CPO) [x1.:230).

'"

OREGANO, Greek - Origanum heracleoticum

Platt' 70: Leaf (TiS) showing.sessile secretory gland on uppersu r face with o i l-fille dsubcunculnr space resulting infull y -e x t e nd ed cu ti cl c .Pallis adc parenchyma visib lebeneath epidermis (Lr-.1. DF.stained with safranin and fastgn:cn)[x 420].

Plat t' 7 1: Sessile sec retoryglan d (T IS) on uppe r lea fsurface with oil-filk-d. fully-extended intact cuticle tLM,DI e. fresh. unstain ed } [ x4::!O].

Plat e 72: Sialked glandulartrichome (TiS) with unicellularhead on upper leaf surface (l~1.OF. stained with safranm andfas t grccm lx4:!OJ.

4J

PEPPERM INT - Mentha piperita

Platt' 7.\: Turgid s..:ssik secretory glands and non-secretory trichome, on calyx surface (cryoSEM )[x 137].

Plai t' 74: Det ai l ofsess ile secretory glands and non -secretory trichornes on calyx surface (cryo-SEM )[:

PERILLA - Perilla frutescens

r illtl' 75: Learn /51showing :! types ofsecretoryglandson lower surface:a) muhiccllular wnh oil-tilled subcu ticularspaceresulting infully-exrendcd cuncleb)stalked with unicetlularbeod (L\1. BF.fresh. uns taincd l[.x2S3].

Plate 76: Leaf IT,S) showing sta lked secretory glands on upper and lower surfaces and single non-secretorytrichome on lower (It\l. BF. fresh. unstain ed] [x2R5].

PLANTA IN - Plantago psyllium

Plate 77: whole seed OIlier soa king inwater fo r severa l hours showingresult an t swolle n. mu c ilagi nou scoaring rl.M . D1 C) lx24 ].

P tar e 78: De tail ed structure ofmuci lagino us coa ling after soaking inwate r for several hourv (S E" !. frl..'C7Cdried )[-'1.57111 .

Plat e 79 : Detail ofmucilaginou-,coatin g (Ti S) attached to seedte sta (SEf\l , f'rcc zc-dri cd j[x266J.

PIal I.' SO: Co ntex tual view ofwhole seed (TIS )and ils mucilaginous layerafter imbib ition (SE~1 . freeze -dr ied] [x4 1].

PLATE 81

Aglls/{/cht'fu('1/;cu/llm at Auchmcruivc herb garden

ROSEMARY - Rosemarinus offic inalis

l'l ure H2: Detai l of freeze..fractured leafto reveal spongy mcsophytt and sec retory glands o n lower surface(c'Y0-SEM ) [1(6 15).

Plat t' 83: Lower surfaccoflcafshowi ng both sess ile and stalked scxrctorv glands together with non-secretoryrrichomcs associated with water conse rvat ion and imparting a gla ucous appearance (cl)'{l-S E\ f) [:'I:2R7].

Pra te Ho4 : Detail of sessilesecretory gland (TIS) showingbasal cell and 4 distinctsecretory cells loosely coveredby cuticle. Epidermis andunderlying p alli s a d cparenchyma clearly visible(LM. BF. stainc-d with safraninand fast greenll 'I( 1.150].

I'l at e 8:5: Dclail of stalkedsec retory gland (TIS) withunicellular head on lower leafsurface (LM. BF, fresh. staine dwith light gr~~n ) [x 1.250].

l'l a" ,' 86: Detai l of sess ilesecretory gland (TISI on lowerleaf surface showing basal celland 4 secretory cells (LM. S F.stained with light green)['1(1.150].

SAGE - Salvia officinalis

TANSY - Tanacetum vulgare

Platt' H7: 1),:13il of scsvilc secretory gland with turgid. intact cuticle on lower lea f surface. Sing fc basal cellvisible (cryo-SEM) [x 1.540] .

Plate loiS: Sessi le secretory glands with turgid . intact cuticles on lower leaf surface. Single dosed stomavisibleIcryo-SEf\1)[x81 2].

PLATE 89

Experimental plot of tansy grown for volatile oil with high thujone content (Auchincruive]

WORMWOOD - Artemisia absinthium

Plate 90: Leaf (TiSI ...hewingsessile secretory gland.. withoil -filled subcuticular spaceresulting in fully-extendedcut ide j lM . DIe. fresh.unstainc-d) [x-416].

I'l al t' 91: Leaf (TiS) showingsessile secretory gland on uppers urface wi t h o i l - fi lledsubcuticular space resulting infully-extended cuticle (U .1.dar k ground illumination. fresh.unstained) [11,416].

I'laic 92: Leaf (TIS) showingsess ile secretory gland on lowersurface with oi l -f i lledsubcuticular ..pan: result ing infully-extended cuticle. :\'00-sec retory tricbomcs clearlyvisible ( L:-'1. DIe. fresh.unstained) [11,416J.

50

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51

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52

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55

PLATE 93

Street market withherbs and spices. Seoul. Korea

LIST OF PLANT SPECIES USED IN AROI\1ATlI ERAPY

I,ATI:\ :\ .\\I E F:;\:GI.ISII :",\\u:Ahics spp. FirAchillea milletolium YarrowAllium sanvum GarlicAllium .\('1)(1 OnionAlpinia J,:a/CIIIKu Gelanual footAmm; 1";.\1/111(1/ Khd laAml"ri.I' balsamiiera AmyrisAn~'tJlllm grovvolens Dil lAnxdictllln:/umgdicd AngelicaAniba msaeodora RosewoodAnthoxanshum of-lorI/ film FjcuveA/,i/lm gran'olt'''1 Cclerv seedArtemisia anntta \\'onnwood sweetArtemisia arborescens Mugwcrt greaterArremisia drucunculus TarragonBenda spp. BirchBosweiiiu curta; FrankincenseBursera delpcvhionu Linaloc bcrrvCmulI/ga odorata Ylane Ylan~Conarium III=on it"/l1T/ E1I:01l-Corum cun"; CarawayCedrus srr. Cedarwo..:)ljChamuemelum spp. Chamomile RomanCinnamomum camphora Camphor('inlla11l01111111/ zevlanicum C innamon leafCi.HlH ladamtems Cistu'>Citrus auranntolia LimeCitrus anrantium Bitter ora nge (JX'i:l lCitrus aunmtinm Ncroli bigaradc (flower]Citrusuunnninm ssp. amara Pctitgrain (kat)

(llIlL~ lIUrtimi/11II ...,p. ht'/galllia Bergamot (pl'dl(i/lln auramium ssp. bergamio Pcutgram bcrgamicrCitrus limol/ Le monCurus parudisi GrapefruitC//IlLI n'liclllala var. .\1t'f1J Wintergreen

lft'lichn~~lIm ,mg/l.~/ili,lill'" Curry planlII/llIllllli \ 11/J!1I111~ flop1(1]!t'riclIlII Ih'l'li,ralll11l SI. John's won

1(IT~ OII/l{ "Jlit-il/ali\' IlyssoplIIicill'" n'n l11/ Anise star1II IIIa I(I'"n'fJ/(,lI~ Inu laJa\l"i~lI/l/I/ ollic'ill\ il-rtll.~ communis MvrtlcXordostachvs jatamansi SpikenardXepeta cataria CatnipOcimum spp_ BasilOriganum spp. OreganoPelargonium spp. GcraniumPesmsetinum sauvum Pars ley seedPicco mariana Spruce (b lack)Pimento dioica Pimento berryPiJII('11fU racemose U3YPimpinella IlIIi.\"/II1I An iseedPinus svlvestris PincPiper cllhena Cubeb sccdPiper nigrum Black pepperPOKOSll'II/0I1pmc/w llli PatchouliPrunus amvgdatus Almo ndPS(,lldof,l"Ilga spr. Fir douglasRavensara aronuuica RavensaraRosa cvntitolia Rose ot toRO.\lI1ar in;ls o{lici l/a!is RosemarySalvia iavanduluctolia Sage Spani shSalvia ollic;lIa{i_\ Sag eSalvia sclaveu ClarySalvia tritoba Sage crctanSantalumaibum San dalwoodSatvrejo hartensis Summer sllvorySaw"('j a lI1o l/fm11l \Vinter sa vorySafl/Ilja thymhra Savory thymc-leavc dSO"l'lH h.,I/:o;1I Benzoin5.\:ygilill/ amll1aliclIlll Cllwe bud

Tagl!f{'.~ glam/li/(/i:ra Tagett:sTlllI;a occillt'lllalis Thuja

ThY"'I/,~ sp p. Thym eTilia t'IIm/hlt'll Lind...n blossomI5l1ga c

GLOSSARY

Aglycoll c the non-carbohydrate port ion ofa glycoside.Azulene cyclopcntacycloheptene [C I,) I ~]. insoluble in water, intensely blue with naphthalene

odour: precursor to chamazulene in plants.Calyx the sepals collectively, forming the outer whorl ofthe flower.Car pup horc the part of a flower axis to which carpels are attached. The extension of the stem

which divides into two and to which each single seed is attac hed (Umbelliferac) .Com missura l face the inner surface ora seed pai r to whic h both seeds arc attached.C rit ical point (dryi ng) the temperature and pressure ofa substance at which the densities of the

liquid and gaseous states are identical. At this point there is no meniscus and artifactscaused by surface tensio n arc avo ided.

Cryogenic fixation a method of mechanical fixation avoiding chemical artifacts where aspec imen is quickly frozen at a very low temperature. usually -2 1O"C in Nitrogen slush andexamined microscopically in the frozcn state.

Cuticle an outer skin or pellicle. sometimes referring to the epidermis as a whole. especiallywhen impcnneab le to water; (bet. I layer of waxy materi al. cutin. on the outer wall ofepide rmal cells in many plants. making them fairly impenneablc to water.

C utin a wax-like or fatty substance impregnating epidermal walls of plant cell s and also forminga separate layer. the cuticle. on the outer wall of the cpidennis in plants making the surfaceimpermeable to water.

Cytop lasm all the living parts ofa cell inside the cell memb rane and excludin g the nucleus.Endocurp the innermost layer of pcricarp of fruit. usually fibrous. hard or stony. as the "stone"

enclosi ng the seed in plums. cherries. etc.Endoplasmic reticulum (E R) extensive. convoluted internal memb rane in cells. continuous

with the outer nuclea r membrane and enclos ing a continuous internal space (lumen).Involved in the synthesis and transport of membrane proteins and lipids and of materialdes tined for secretion from the cell.

Epithelium sheet of cells tightly bound toget her. lining any internal or external surface inmulticellular organisms. Epithelia variously serve protecti ve, secretory or absorptivefunctions.

Esscnrlal oils mixtures of various volatile oils de rived from benzenes and tcrpcncs found inplants and producing characte ristic odours. Having various functions such as attractinginsects or \...-arding offfungal attacks.

Flavedo the outer layer. or rind. ofpericarp in citrus fruits.G lycoside any class ofcompound whichon hydro lysis gives a sugarand a non-sugar residue.Isupentenyl pyrophosphate biogenetic precursor ofterpcnes [c'H.].Lacuna space or cavity,Leucoplastt id) colourless plastid that develops into chloroplasts, etc.Llg nin a hard material found in walls of cells of xylem in plants. a very variable cross-linked

polymer which stiffens the cell wall.Lipophilic relating to or having a strong affinity for fats and other lipids.

,"

Lumen internal space ofany tubular or sac-like organ or sub-cellular organelle.l\lesocarp the middle layer of the pericarp e.g. comprising the flesh of fruits such as plums and

cherries.l\litochondria organell es in the cytoplasm of eukaryotic cells, having a double membrane,

important for energy production and genetic coding.Parenchyma soft plant tissue composed of thin-walled , relati vely undifferentiated cells which

may vary in structure and function .Pericarp the tissues of fruit that develop from the ovary wall, comprising an outer skin,

sometimes a fleshy mcsocarp and an innerendocarp.Perisperm in some seeds , a storage tissue formed by proliferation of the nucleus rather than the

endosperm.Phloem the principal food-co nducting tissue of vascular plants extending throughout the plant

body. It is composed of elongated conducting vessels and sieve tubes (in angiosperms],both containing clusters of pores (sieve areas) in the walls through which the protoplasts ofadjace nt cells communicate. Sugars and amino acids arc the main nutrients transported viathe phloem. Parenchymatous companion cells closely assoc iated with the conductingelements are involved in the del ivery to- and uptake of- material from the phloem.

Plasmodesmata cytoplasmic threads runnin g transversely through plant cell walls andconnecting cytoplasm ofadjacent cells.

Plastid one of a class of cytoplasmic organelles, includin g chloroplasts, found in plants andcukaryotic algae. Bounded by a double membrane and containing DNA.

Proliferation increase by frequent and repeated reproduction ; increase by cell division.Protoplast the living component ofa cell , i.c. the protoplasm not including any cell wall.Rhizome thick horizontal stem, usually underground , bearing buds and scale leaves and sending

out shoots from above and below.Secondary metabolites compounds produ ced by plants and microbes, c.g. antibiotics, alkaloids.

and flower pigments that are not essential to the growth ofthe organism.Secretory tissues cells and tissues that secrete substances such as digestive enzymes.

polypeptide horrnones, neuro-transmitters or complex material such as mucus, slime.fragrances , etc.

Senescence advancing age; the complex ageing processes that eventually lead to death.Thujone [CIIJH1"O] a toxic constituent of many essential oils. A colourless liquid practically

insoluble in water.Tracheid a type of water-conducting xylem cell present in all vascular plants with lignified

seco ndary cell wall usually conta ining spiral thickening or bordered pits.Trichome any of various outgrowths of the epidermis in plants including branched and

unbranched hairs, vesicles, hooks, spines and stinging hairs; a hair tuft .Xylem the main water-conducting tissue in vascular plantsextending throughout the whole plant,

involved in storage, support and the transport of minerals and nutr ients. Xylem iscomposed oftracheary elements: tracheids and (in angiosperms) vessel elements. Both areelongated. hollow cells with thickened, usually heavily-lignified wa lls and lackingprotoplasts when mature. They are joined end-to-end to form a continuous, conductingtube.

5"

INDEX OF SPECI ESE:>G Ll SII :>'\~ I ES LATI:> :>A~IES

Page no: I a~e no:

Basil 15Bog myrt le 16

Caraway 17Catmint 18Cedar 19C hamomile, Roman 20

Citrus 21Clove .22Cumin 23Dragonhead 24

Euca lyptus 26Frankincense 27Ginger 30Hyssop 3 1

Juniper 32Korean mint 33

Lavender 34Lemon balm 35Lovage 36l\..1arjoram 37Myrrh 28Nutmeg 38Orchid. lady's slipper 39

Oregano 40Oregano. Greek .4 1Peppermin t 42

Perilla ..43Planta in 44Rosemary 46Sage ..47Tansy 48

Wormwood 50

Agastache rugosa 33Artemisia absinthium 50

Boswellia carteri 27

Carum carvi 17Cedru s spp 19Chomaemelum nobile 20

Citrus spp 2 1

Commiphora myrrha 28

Cuminum cyminum 23Cy pripedium pubescens var; catceolus 39Dracocephalum moldavica 24Eucalyptus citriodora 26

Hvssopus officinalis 3 1Juniperus communis 32Lavandulu angustifotia 34l.evisticum officinale 36Melissa officinalis 35Mentha piperita .42Myrica ga le 16M yristica fragrans 38Nepeta cataria var. citriodora 18Ocimum bas ilicum 15Origanum marj orana 37Origanum vulgare .40Origanum heracleoticum .4 1Perilla frutescens .43Plantago psyllium -14Rosemarinns officinalis _ .46Salvia officinalis .47Syzygium aromaticum 22

Tanacelllm vulgare ,48Zingiber officinate 30

.. ,

PLATE 94

Statue of a SlrCCI seller with various herbs essent ial oils and perfumes(Perfumery Museum, GmsI:. South of France)

ISBN 0-9538461-0-5

j ~llJjn lUl,llljl~.=::: .--1

,

Dr. K. P. Svoboda lectures on plant anatomy, plant breeding andbiotechnology at the Scottish Agricultural College (SAC), Auchincruive.Slie is also responsible for research on aromatic and m dicinal plantsIncludfnq botany, hysiology, chemotaxonomy and the production ofsecondliry metabolites and theirapplication. During the est 10years shehas published 73 refereed and general articles on aromatic andmedicinal plants and has contributed over 70 papers and posters tonational and international conference. She has travelled extensively intheco rseofherwork and in1992 spentayearinJapan. ,.

'"

TOMAS GALEN SVOBODAT. G. Svoboda isa first-yea studentat MedicarUniverslty,Dundee, He hasan interest in aromatic plants nd essential oils and has worked at a -chamomile far in Norfolk running a larg -scele distillation plant herH has now finished his project t SAC on the pli}'!ochemicainv stigation and genetic fingerprinting of ernomlle.: nd during thistime also contributed tosev~ral papers onvarious otheraromatrc plants.

f Y , .

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