Glandular trichomes, secretory tissues of  plant volatile compounds

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. . . Trichomes. . Trichomes. . Plant hairsEpidermal appendages of diverse form, structure, and functionsDistinguished from emergences. . Morphological categories. . PapillaeMulticellular trichomesTwo- to five-armed trichomesStellate trichomesPeltate trichomesBranched trichomesRoot hairs. .

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Glandular trichomes, secretory tissues of plant volatile compounds

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1. April 24, 2008 Jeong Hwa Song Vegetable Science Lab Seoul National University Glandular trichomes, secretory tissues of plant volatile compounds ?? ?? ?? ?? ??? ?? ?????? glandular trichome? ??? ???????. ?? ???? ??? ??? ??? ? ?? ??? ? ?????, ?? trichome? ???? ???? ? ? ???? ???? ???. ?? ?? ?? ?? ??? ?? ?????? glandular trichome? ??? ???????. ?? ???? ??? ??? ??? ? ?? ??? ? ?????, ?? trichome? ???? ???? ? ? ???? ???? ???.

3. Trichomes Plant hairs Epidermal appendages of diverse form, structure, and functions Distinguished from emergences Trichome?? ??? ??? ???? ????, ??? ??, ?? ? ??? ???. ?? emergence? ????? ???, ??? emergence? ??? ?? ????, ?? ?? ?????? ??? ??? Trichome?? ?? ??? ??? ?????. Trichome?? ??? ??? ???? ????, ??? ??, ?? ? ??? ???. ?? emergence? ????? ???, ??? emergence? ??? ?? ????, ?? ?? ?????? ??? ??? Trichome?? ?? ??? ??? ?????.

4. Morphological categories Papillae Multicellular trichomes Two- to five-armed trichomes Stellate trichomes Peltate trichomes Branched trichomes Root hairs Trichome? ? ??? ?? 7?? ????? ??? ? ????. ????? ?? ??? ??? ??, ???? ???? ?? multicellular trichome ?? 2??? 5? ?? ??? ?? two-to five armed trichome ? ??? ??? ?? stellate trichome ?? ??? head cell? ?? peltate trichome ??? ?? branched trichome ??? ?? ????. ? ? ?????? ??? ??? multicellular, peltate trichome ?? ??? glandular trichome? ?????. Trichome? ? ??? ?? 7?? ????? ??? ? ????. ????? ?? ??? ??? ??, ???? ???? ?? multicellular trichome ?? 2??? 5? ?? ??? ?? two-to five armed trichome ? ??? ??? ?? stellate trichome ?? ??? head cell? ?? peltate trichome ??? ?? branched trichome ??? ?? ????. ? ? ?????? ??? ??? multicellular, peltate trichome ?? ??? glandular trichome? ?????.

6. Glandular trichomes glandular trichome? ???? ??? ??? ????. ??????? ??? basal cell, stalk cell, secretory cell ? ??? ??? ???? ???? oil storage cavity? ?????. ?? cell? ???, ??? ??, ???? ?? ???? ?????, ??? ?? ???? cell ??? ???? ????? ?? ?? ??? ??? ?? ?? ??? cavity? ???? ???. glandular trichome? ???? ??? ??? ????. ??????? ??? basal cell, stalk cell, secretory cell ? ??? ??? ???? ???? oil storage cavity? ?????. ?? cell? ???, ??? ??, ???? ?? ???? ?????, ??? ?? ???? cell ??? ???? ????? ?? ?? ??? ??? ?? ?? ??? cavity? ???? ???.

7. Glandular trichomes ???? Glandular trichome? ????? ??????. ??? ??? ?? ????? ???? ???? basal cell, stalk cell, head cell? ???? ?? Head cell? ?? ??? ???? ????? Basal cell stalk cell? ?? ??? ???. head cell ??? ?? essential oil? ?? ? ??? ?????. ??? ???? ????? ???? cavity ? ?? ?? ??, ? ?? ??? ???? ???? ??? ????. ??? ???? head cell? stalk cell? ???? basal cell? ? ??? ????. Glandular trichome? ????? ??????. ??? ??? ?? ????? ???? ???? basal cell, stalk cell, head cell? ???? ?? Head cell? ?? ??? ???? ????? Basal cell stalk cell? ?? ??? ???. head cell ??? ?? essential oil? ?? ? ??? ?????. ??? ???? ????? ???? cavity ? ?? ?? ??, ? ?? ??? ???? ???? ??? ????. ??? ???? head cell? stalk cell? ???? basal cell? ? ??? ????.

8. Functions of glandular trichomes Contributing to the flavor and aroma Attracting pollinators Aiding in the dispersal of certain fruits Protecting from herbivore attack Exuding lipophilic esters, terpenoids, waxes, flavonoid aglycones ?? ??? ?? ???? ??????? ?????? ?????? ???? ??? ?? ????? ???. ????? ?????? ???? ??? ???. ?????? ???? ???????, ???, ???, ?????? ???? ?? ?????. ?? ??? ?? ???? ??????? ?????? ?????? ???? ??? ?? ????? ???. ????? ?????? ???? ??? ???. ?????? ???? ???????, ???, ???, ?????? ???? ?? ?????.

9. Glandular trichome? ?? ????? ?? ???? ?????. ??A? ??? ?? ??? glandular trichome?? B? brushing? ?? trichome? ??? ?????. ?? ?? ?? ??? DVT? ??? ? ??, Epidermal strip?? midvein section ?? brush ??? ? ? ??? ???? ???? ?? ? ? ????. ?, DVT?? diterpene?? trichome head??? ?? ? ???? ?? ? ? ????. DVT=duvatriene-1,3 diols (diterpene) Glandular trichome? ?? ????? ?? ???? ?????. ??A? ??? ?? ??? glandular trichome?? B? brushing? ?? trichome? ??? ?????. ?? ?? ?? ??? DVT? ??? ? ??, Epidermal strip?? midvein section ?? brush ??? ? ? ??? ???? ???? ?? ? ? ????. ?, DVT?? diterpene?? trichome head??? ?? ? ???? ?? ? ? ????. DVT=duvatriene-1,3 diols (diterpene)

10. Glandular trichomes as secretory tissues ? ??? ?? ??? ?? glandular trichome? ???? ?, ??? ?? ?? ??? ????. ??? ? ??? ?? ????? HPLC? ???? ??? ???, ?????? blank, sesquiterpene?? artemisitene, artemisinin? ????, ??? ????? ?? ?? essential oil? ??????. ????? ???????, glandular trichome? ???? ???? ?? ??? ???? glandular trichome? ?? ???? ??? ?? ??? ??? ?????. ?? ?? sesquiterpene?? ?? glandular trichome???? ????? ?? ? ? ????. ? ??? ?? ??? ?? glandular trichome? ???? ?, ??? ?? ?? ??? ????. ??? ? ??? ?? ????? HPLC? ???? ??? ???, ?????? blank, sesquiterpene?? artemisitene, artemisinin? ????, ??? ????? ?? ?? essential oil? ??????. ????? ???????, glandular trichome? ???? ???? ?? ??? ???? glandular trichome? ?? ???? ??? ?? ??? ??? ?????. ?? ?? sesquiterpene?? ?? glandular trichome???? ????? ?? ? ? ????.

11. ????? ?? ??? trichome? ??? ?? ?????, ?? ?? ???? ???????. ????? ?? ??? trichome? ??? ?? ?????, ?? ?? ???? ???????.

12. Secretion capacity related to structure type Main types of glandular trichome in tomato types IV and VI Trichome ??? ??? ??? ?? ?? ????? ??? ? ????. ?? ?????? 7?? ??? trichome type? ???? ???, ??? ?? ??? ???????. ? ? glandular trichome? main type? type 4? 6?? ?? ???? ??????. ??? type? ?? ????? ?????, 6? ???? ??????? ??? glucose ester?? ??? 4? ???? ??? ????? ?????. Trichome ??? ??? ??? ?? ?? ????? ??? ? ????. ?? ?????? 7?? ??? trichome type? ???? ???, ??? ?? ??? ???????. ? ? glandular trichome? main type? type 4? 6?? ?? ???? ??????. ??? type? ?? ????? ?????, 6? ???? ??????? ??? glucose ester?? ??? 4? ???? ??? ????? ?????.

13. Trichome type and secretion ?? ?? ??? ?????? ??? ?? ??? ????? ???? sesquiterpene? ??? zingiberene? trichome type? ?? ??? ??? ????. ??? ??? zingiberene? ?? ?????, ?? ?? type 4? ???? ? ??? ???? ?? ? ? ????. ??? type 6? non=glandular type? ???? ??? ??? ??? ?? ????. ?? ?? ??? ?????? ??? ?? ??? ????? ???? sesquiterpene? ??? zingiberene? trichome type? ?? ??? ??? ????. ??? ??? zingiberene? ?? ?????, ?? ?? type 4? ???? ? ??? ???? ?? ? ? ????. ??? type 6? non=glandular type? ???? ??? ??? ??? ?? ????.

14. Trichome type and secretion ?? ?? ??? ????? ????. Zingiberene? type 4? ????? ??? ? glandular type?? ???? ?? ???, Type6? non-glandular type?? ????? ????. ?? sesquiterpene? zingiberene? type 4???? ???? ?? ? ? ????. ?? ?? ??? ????? ????. Zingiberene? type 4? ????? ??? ? glandular type?? ???? ?? ???, Type6? non-glandular type?? ????? ????. ?? sesquiterpene? zingiberene? type 4???? ???? ?? ? ? ????.

15. Secretion capacity related to photosynthesis Non-photosynthetic glandular trichomes Sucrose to the monoterpenes and sesquiterpenes Peppermint Photosynthetic glandular trichomes Carbone dioxide to diterpenes Tomato, tobacco, sunflower, and geranium ?? ??? ?? ??? trichome? ??? ??? ???? ??? ? ????. ???? ?? ?? trichome? ???? sucrose? ?? ???? terpene ???? ????? ???? monoterpenes and sesquiterpenes? ?? ? ?????. ???? ????? ????? ???? ?? glandular trichomes? ?????? ???? ?? monoterpenes and sesquiterpenes ? ?? diterpene? ?? ? ?????. ?? ???? ??? ???, ??, sunflower, ??????. ?? ??? ?? ??? trichome? ??? ??? ???? ??? ? ????. ???? ?? ?? trichome? ???? sucrose? ?? ???? terpene ???? ????? ???? monoterpenes and sesquiterpenes? ?? ? ?????. ???? ????? ????? ???? ?? glandular trichomes? ?????? ???? ?? monoterpenes and sesquiterpenes ? ?? diterpene? ?? ? ?????. ?? ???? ??? ???, ??, sunflower, ??????.

16. Photosythesis capacity and secretion ?? ???? ?? ??? ?? ????? DVT? ?? ?? ??? ??? ????. ?? ?? ??? ???? ?? ??? ???? ???, ?? ?? ???? 97% ?????. ??, ????? ?????? ???? ??? DCMU HgCl2? ???? ??? DVT ???? ?? ??? ????? ??????? ??? ???? ??? ???? ??? ???? ?????. ????? ????? sucrose? ??? ? 34% ??? ??? ??? ?? ??? ??? ???? diterpene? ????? ?? ? ? ????. DCMU, HgCl2-???? ????? ?? ?? Antimycin – ??????? ????? ?? ?? Photosynthetic DVT=duvatriene-1,3 diols (diterpene)?? ???? ?? ??? ?? ????? DVT? ?? ?? ??? ??? ????. ?? ?? ??? ???? ?? ??? ???? ???, ?? ?? ???? 97% ?????. ??, ????? ?????? ???? ??? DCMU HgCl2? ???? ??? DVT ???? ?? ??? ????? ??????? ??? ???? ??? ???? ??? ???? ?????. ????? ????? sucrose? ??? ? 34% ??? ??? ??? ?? ??? ??? ???? diterpene? ????? ?? ? ? ????. DCMU, HgCl2-???? ????? ?? ?? Antimycin – ??????? ????? ?? ?? Photosynthetic DVT=duvatriene-1,3 diols (diterpene)

17. Photosythesis capacity and secretion ??? ?????? GPP? ????? ?? ??? ????? ? monoterpene? ??? ??? ????. ????? GPP? ???? ???? monoterpene ???? ???? ???? limonene? ??? ??? NAD ??? ???? ??? ???? ?? ? ??? ??? menthone ? ??? ????? ?? glucose-6-phosphate? ??? ??, menthone? ??? ??? ?????, ?? G6P? ???? ???? ??? menthone? ???? ?? ?????. ?? G6P ??? ???? cibacrone blue? ??? ???? ??? ???? ?? ???? ?????. ?, ????? ??? tichome?? ????? sucrose? ??????? monoterpene?? ??? ????? ?? ?????. ??? ?????? GPP? ????? ?? ??? ????? ? monoterpene? ??? ??? ????. ????? GPP? ???? ???? monoterpene ???? ???? ???? limonene? ??? ??? NAD ??? ???? ??? ???? ?? ? ??? ??? menthone ? ??? ????? ?? glucose-6-phosphate? ??? ??, menthone? ??? ??? ?????, ?? G6P? ???? ???? ??? menthone? ???? ?? ?????. ?? G6P ??? ???? cibacrone blue? ??? ???? ??? ???? ?? ???? ?????. ?, ????? ??? tichome?? ????? sucrose? ??????? monoterpene?? ??? ????? ?? ?????.

18. ????? ?? ??? ?? ???? ??? ??? ????. ????? ?? ??? ?? ???? ??? ??? ????.

19. Secretion capacity related to the environment Growth environment Light Temperature Water Chemicals Salts Heavy metals Hormones Physical damage Insect damage ? ???? ?, ??, ??? ?? ??? ????, ????? ?, ??? ?? ?? ????? ??, ??? ?? ??? ?? ?? ???, ?? ??? ? ??? ?? ??????. ? ???? ?, ??, ??? ?? ??? ????, ????? ?, ??? ?? ?? ????? ??, ??? ?? ??? ?? ?? ???, ?? ??? ? ??? ?? ??????.

20. Secretion capacity related to the environment - light ??, ??? ??? ?? ?? ? ???? ?? ??? ??? tichome? ??? ??? ?????. ?? ???? ??? ????? ??? ???? ?? ???? ??? ??? ? ? ????. ??? ????? trichome ??? ?????, ?? ???? ??? ?? ? ? ????. ??? ?? ??? trichome ??? ??? ??? ?? ? ? ????. ??, ??? ??? ?? ?? ? ???? ?? ??? ??? tichome? ??? ??? ?????. ?? ???? ??? ????? ??? ???? ?? ???? ??? ??? ? ? ????. ??? ????? trichome ??? ?????, ?? ???? ??? ?? ? ? ????. ??? ?? ??? trichome ??? ??? ??? ?? ? ? ????.

21. Secretion capacity related to the environment - water ??? ?? ????? ?? ??? glandular trichome? ??? ??? ????. ?? ????? ??? ? apical zone?? basal zone ???? trichome ??? ???? ? ? ????. ?? trichome cell? ??? ???? ??? ???? ???? ?????. ??? ?? ????? ?? ??? glandular trichome? ??? ??? ????. ?? ????? ??? ? apical zone?? basal zone ???? trichome ??? ???? ? ? ????. ?? trichome cell? ??? ???? ??? ???? ???? ?????.

22. Secretion capacity related to the environment - Salts ??? ??? ??? ? stress? ??? ? trichome? ?????. Stress? ??? head cell?? ????? ????, stalk cell? ?????? ?????. ?? ?? ?? ??? ??? ??? ?? ??? ?????. A? ? ????, B? head?? ??? ??????, C? ??? stalk cell?? ???????. ? ??? ?????? ? ??????? ???? ? ??? ??? ?? ?????. ?? ??? ? ??? ?? ?????? ?? ????, ????? ??? ???? ??? ?? ?? ??? ???????. ???? stalk cell ??? Na Cl ??? ?? ?? ? ? ????. ?, ??? ? ????? ??? NaCl? ???????? ?????? ??? ??? ?? ? ? ????. ?? ?? ????? ???? ??? ?????? trichome? ???? ?? ??? ????? ?? ? ? ????. 0.05M NaCl EDX-energy dispersive x-ray analysis??? ??? ??? ? stress? ??? ? trichome? ?????. Stress? ??? head cell?? ????? ????, stalk cell? ?????? ?????. ?? ?? ?? ??? ??? ??? ?? ??? ?????. A? ? ????, B? head?? ??? ??????, C? ??? stalk cell?? ???????. ? ??? ?????? ? ??????? ???? ? ??? ??? ?? ?????. ?? ??? ? ??? ?? ?????? ?? ????, ????? ??? ???? ??? ?? ?? ??? ???????. ???? stalk cell ??? Na Cl ??? ?? ?? ? ? ????. ?, ??? ? ????? ??? NaCl? ???????? ?????? ??? ??? ?? ? ? ????. ?? ?? ????? ???? ??? ?????? trichome? ???? ?? ??? ????? ?? ? ? ????. 0.05M NaCl EDX-energy dispersive x-ray analysis

23. Secretion capacity related to the environment – physical damage ??? ??? ??? Datura?? ??? ?? ??? damage? ??? ? ?? ?? ??? b-caryophyllene? ??? ?? ? ? ?? ??? ??? ??? ?????. ??? ???? ?? ???? ??? ? –caryophyllene (sesquiterpene)? ??? ?????, ? ????? ?? ??????. ?? ??? ???? ?? ????? ?? terpene??? ????? ?????. ??? ??? ??? Datura?? ??? ?? ??? damage? ??? ? ?? ?? ??? b-caryophyllene? ??? ?? ? ? ?? ??? ??? ??? ?????. ??? ???? ?? ???? ??? ? –caryophyllene (sesquiterpene)? ??? ?????, ? ????? ?? ??????. ?? ??? ???? ?? ????? ?? terpene??? ????? ?????.

24. Summary and further study Most plant volatile compounds are derived from glandular trichomes. Secretion capacity is related to the trichome type and photosynthesis capacity. Secretion capacity is related to the plant growth environment, chemicals, and physical damage. There are few studies about relations among plant growth environment, trichome structure, and plant volatile compounds.

25. References Bosabalidis, A.M.and F. Exarchou. 1995. Effect of NAA and GA3 on leaves and glandular trichomes of Origanum x intercedens Rech. International journal of plant sciences. 156: 488-495. Choi, Y.E., E. Harada, G.H. Kim, E.S. Yoon, and H. Sano. 2004. Distribution of elements on tobacco trichomes and leaves under cadmium and sodium stresses. Journal of Plant Biology. 47: 75-82. Duke, M.V., R.N. Paul, H.N. Elsohly, G. Sturtz, and S.O. Duke. 1994. Localization of Artemisinin and Artemisitene in Foliar Tissues of Glanded and Glandless Biotypes of Artemisia annua L. International journal of plant sciences. 155: 365-372. Esau, K. 2006. Esau's Plant Anatomy. John Wiley & Sons New Jersey. Hare, J.D. 2007. Variation in herbivore and methyl jasmonate-induced volatiles among genetic lines of Datura wrightii. Journal of Chemical Ecology. 33: 2028-2043. Keene, C.K.and G.J. Wagner. 1985. Direct Demonstration of Duvatrienediol Biosynthesis in Glandular Heads of Tobacco Trichomes. Plant Physiol. 79: 1026-1032. Maluf, W., G. Campos, and M. das Graças Cardoso. 2001. Relationships between trichome types and spider mite (Tetranychus evansi) repellence in tomatoes with respect to foliar zingiberene contents. Euphytica. 121: 73-80. McCaskill, D.and R. Croteau. 1995. Monoterpene and sesquiterpene biosynthesis in glandular trichomes of peppermint (Mentha x piperita) rely exclusively on plastid-derived isopentenyl diphosphate. Planta. 197: 49-56. McCaskill, D.and R. Croteau. 1999. Strategies for bioengineering the development and metabolism of glandular tissues in plants. Nature. 17: 31-36. Nihoul, P. 1993. Do light intensity, temperature and photoperiod affect the entrapment of mites on glandular hairs of cultivated tomatoes? Experimental and Applied Acarology. 17: 709-718. Sam, O., E. Jeréz, J. Dell'Amico, and M.C. Ruiz-Sanchez. 2000. Water Stress Induced Changes in Anatomy of Tomato Leaf Epidermes. Biologia Plantarum. 43: 275-277. Turner, G., J. Gershenzon, E.E. Nielson, J.E. Froehlich, and R. Croteau. 1999. Limonene synthase, the enzyme responsible for monoterpene biosynthesis in peppermint, is localized to leucoplasts of oil gland secretory cells. Plant Physiology. 120: 879-886.

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