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The document describes the comparative histoanatomical aspects of the fruits of five Apiaceae species - dill, cumin, carrot, fennel, and anise. It finds that the fruits have the same basic structural organization of epicarp, mesocarp and endocarp layers closely united with the seed. The epicarp is one cell layer thick with isodiametric cells in all species. The mesocarp is multiple cell layers thick and parenchymatous, with some differences in cell size and number of layers between species. Vascular bundles are present in the mesocarp and secretory ducts vary in size and number between species. The fruits show some differences in shape and cell features but share the
The document describes the comparative histoanatomical aspects of the fruits of five Apiaceae species - dill, cumin, carrot, fennel, and anise. It finds that the fruits have the same basic structural organization of epicarp, mesocarp and endocarp layers closely united with the seed. The epicarp is one cell layer thick with isodiametric cells in all species. The mesocarp is multiple cell layers thick and parenchymatous, with some differences in cell size and number of layers between species. Vascular bundles are present in the mesocarp and secretory ducts vary in size and number between species. The fruits show some differences in shape and cell features but share the
The document describes the comparative histoanatomical aspects of the fruits of five Apiaceae species - dill, cumin, carrot, fennel, and anise. It finds that the fruits have the same basic structural organization of epicarp, mesocarp and endocarp layers closely united with the seed. The epicarp is one cell layer thick with isodiametric cells in all species. The mesocarp is multiple cell layers thick and parenchymatous, with some differences in cell size and number of layers between species. Vascular bundles are present in the mesocarp and secretory ducts vary in size and number between species. The fruits show some differences in shape and cell features but share the
265 COMPARATIVE HISTOANATOMICAL ASPECTS OF THE FRUIT OF SOME APIACEAE LINDL. FRUIT USED FOR THERAPEUTIC PURPOSES
Rodica Bercu, Livia Broasc
FACULTY OF NATURAL AND AGRICULTURAL SCIENCES OVIDIUS UNIVERSITY, CONSTANTZA, ROUMANIA
Summary The present paper discloses histoanatomical aspects, comparatively to some Apiaceae species fruits such as Anethum graveolens L. (dill), Carum carvi L. (cumin), Daucus carota L. subsp. sativus (Hoffm) Arc. (carrot), Foeniculum vulgare Mill. (fennel), and Pimpinela anisum L. (anise), all of them considered food value but are used for therapeutic purposes as well. The fruits, of all studied Apiaceae species have the same structural plan of organization (epicarp, mesocarp and endocarp), all fruit parts being closely united among themselves and with the seed. Epicarp is one-layered with isodiametric cells (all species), mesocarp is many-layered and parenchimatous in nature (all species), differences arise in terms of number of layers and size of cells that compose it. In mesocarp the fruits of all 5 studied species possess pore developed vascular bundles in the ribs being more developed to cumin and fennel. Secretive ducts, present in mesocarp, are similar in all studied species differences arise in particular as regards their size (the larger in cumin and the smaller in anise fruit) and their number (30-31 in anise fruit and 6 the other species). The mechanical tissue is represnted by few sclerenchymatous elements, accompanying the vascular bundles. The fruit of carrot possesses emerging (thorns) and simple unicellular protective hairs, the latter present in the fruit of anise as well. All studied species are considered food value but are used for therapeutic purposes as well. Key words: histoanatomy, Anethum graveolens, Carum carvi, Foeniculum vulgare, Daucus carota. subsp. sativus, Pimpinela anisum
rodicabercu@yahoo.com; broascalivia@yahoo.com
Introduction The Apiaceae or Umbelliferae, commonly known as carrot or parsley family, is a group of mostly aromatic plants with hollow stems (French, 1971), fruits are non- fleshy schizocarp of two mericarps, each with a single seed; they separate at maturity and are dispersed by wind. Some fruit segments (like those in Daucus ssp. are covered in bristles and spread via external transport. The seeds have an oily endosperm (Watson & Dallwitz, 2011) and generally contain large quantities of fatty oils, with the fatty acid petroselinic acid occurring universally throughout the family while rarely being found outside of the Apiaceae (She et al., 2005). The family is large, with more than 3,700 species spread across 434 genera, it is the sixteenth largest family of flowering plants (Stevens, 2008). Anethum graveolens L. originated in the Mediterranean regions, southern central and southern Asia. Nowadays it is cultivated throughout Europe and North and South America. Dill is a perenial plant, it has a tap root like a carrot, with one long, hollow stalk coming from the root. The dill plant grows up to 75 cm. It has a feathery stalk which produces umbels of tiny yellow flowers. The seeds are numerous, they are oval, about 1/6" long, flattish with five ribs, lightweight and pungent. Fruits contain volatile oil (carvone and limonene), are used for colic, also a combination of dill and fennel, both herbs contain stomach-soothing oils. Carum carvi L. (Caraway) also known as meridian fennel or Persian cumin is a biennial plant, native to western Asia, Europe Annals of RSCB Vol. XVII, Issue 1/2012
266 and Northern Africa. The plant is similar in appearance to a carrot plant, with finely divided, feathery leaves with thread-like divisions, growing on 2030 cm stems. The main flower stem is 4060 cm tall, with small white or pink flowers in umbels. Caraway fruits (erroneously called seeds) are crescent- shaped achenes, around 2 mm long, with five pale ridges. The fruits contain essential oils (carvone, limonene, etc.), it has carminative effect, also is used in digestive problems. Daucus carota subsp. sativus is a flowering plant, native to temperate regions of Europe, southwest Asia and naturalized to northeast North America and Australia; domesticated carrots are cultivars of a subspecies, Daucus carota subsp. sativus. Daucus carota is a variable biennial plant, usually growing up to 1 m tall and flowering from June to August. The umbels are claret- colored or pale pink before they open, then bright white and rounded when in full flower, measuring 37 cm wide with a festoon of bracts beneath; finally, as they turn to seed, they contract and become concave like a bird's nest. The dried umbels detach from the plant, becoming tumbleweeds. Similar in appearance to the deadly poison hemlock, Daucus carota is distinguished by a mix of bi-pinnate and tri- pinnate leaves, fine hairs on its stems and leaves, a root that smells like carrots, and occasionally a single dark red flower in its center. The fruits used in hair and skin problems, also contain essential oils. Foeniculum vulgare is a perennial herb. It is erect, glaucous green, and grows to heights of up to 2.5 m, with hollow stems. The leaves grow up to 40 cm long; they are finely dissected, with the ultimate segments filiform (threadlike), about 0.5 mm wide. Its leaves are similar to those of dill, but thinner. The flowers are produced in terminal compound umbels 5-15 cm wide, each umbel section having 20-50 tiny yellow flowers on short pedicels. The fruit is a dry seed from 4-10 mm long, half as wide or less, and grooved. The fruits comes contain anethole, polymers, volatile oils, etc. Carminative properties of fennel are known from ancient times. Pimpinela anisum also called aniseed is a flowering plant, native to the eastern Mediterranean region and Southwest. Anise is an herbaceous annual plant growing to 0.91 m tall. The leaves at the base of the plant are simple, 3-5.1 cm long and shallowly lobed, while leaves higher on the stems are feathery pinnate, divided into numerous leaves. The flowers are white, approximately 3 mm diameter, produced in dense umbels. The fruit is an oblong dry schizocarp, 3-5 mm long. It is these seed pods that are referred to as "aniseed". Is one of the earliest aromatics mentioned in literature (Stern, 1994). The main use of anise was for its carminative and pectoral effect, like fennel, contains anethole, a phytoestrogen, volatile oil and has been used to treat menstrual cramps (Albert-Puleo, 1980). Little information is available about fruit anatomy of studied species, while the comparative studies lack almost entirely. The present paper on fruit features of this 5 species may contribute to the better knowledge of this group of plants.
Material and methods The mature fruits were collected from field crops of the studied species and were fixed in FAA (formalin:glacial acetic acid:alcohol 5:5:90). For the anatomical study, freehand sections were made (Bercu & Jianu 2003). The samples were stained with alumcarmine and iodine green. Anatomical observations and micrographs were performed with a BIOROMT brightfield microscope, equipped with a TOPICA 6001A video camera. The microphotographs were obtained from the video camera through a computer.
Results and discussion The fruit shape in cross sections is different from some of the species. Thus, it is trapezoidal-coasted like (dill, carrot and fennel) (Fig. 1, A, C; Fig. 2, A), with three projections main ribs, and two lateral (lateral wings), nearly circular-coasted like (cumin) with five ribs, 3 on the dorsal and
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Fig. 1. Cross sections of the fruit of Anethum graveolens (A), Carum carvi (B) and Foeniculum vulgare (C) - ensemble: Ed- endocarp, Ep- epicarp, LW- lateral wings, Ms- mesocarp, S- seed, SD- secretive ducts, VB- vascular bundle.
Fig. 2. Fruit of Daucus carota subsp. sativus (A), and Pimpinela anisum (B) - ensemble observed in cross sections: LW- lateral wings, S- seed, Scl- sclerenchyma, SD- secretive ducts, VB- vascular bundle.
A
A
B
B
C VB S S S
LW
LW
SD
Ep
Ed
Ms VB
LW
SD
Scl S S
SD
VB
x 46
x 43
x 35
x 55
x 35 Annals of RSCB Vol. XVII, Issue 1/2012
268 2 to the underside (Fig. 1, B). The mericarps are almost kidney-shaped to anise due to the reduced pericarp (Fig. 2, B). The pericarp (epicarp, mezocarp and endocarp) (Andrei & Predan, 2001) is closely linked together (all species). The unistratose epicarp consists of almost isodiametric cells, slightly tangentially elongated, with straight side walls (all species).
Fig. 3. Cross sections of the fruit of Anethum graveolens (A), Carum carvi (B) and Foeniculum vulgare (C) - details: Es- endosperm, Scl- sclerenchyma, SD- secretive ducts, T- tegument, VB- vascular bundle.
Fig. 4. Fruit of Daucus carota subsp. sativus (A), and Pimpinela anisum (B) in cross section - details: PH- protective hairs, SD- secretive ducts, Th- thorn, VB- vascular bundle.
The many-layered mesocarp is parenchymatous in nature, with large cells, and thin cellulosic walls. In the mezocarp right main ribs, the vascular collateral bundles are present. In valecule are present secreting elongated and septum ducts as well (all species) (Batanouny, 1992) (Fig. 3, A-C, Fig. 4, A, B). The ventral face contour (adaxial) of the dill fruit is slightly sinuous, almost plane and waved those of the carrot, fennel and cumin. Adaxial vascular bundles and either the two secretive ducts are present
A
A
B
B
C
Scl
VB
SD
SD
Es
T
SD
SD
PH
PH
x 95
x 85
x 117
x 125
x 330
VB
Th Annals of RSCB Vol. XVII, Issue 1/2012
269 (all species). Adaxial are alternating secretive ducts with as many vascular bundles in all fruits mesocarps. In the lateral wings, the dill and fennel fruit vascular bundles are more developed than the rest. Those of carrot wings possess thorns (Fig. 4, A), with sclerenchymatous basal elements. The cumin fruit vascular bundles are placed in the 5 ribs but the valecule secreting ducts are larger and slightly elongated, located one in a latero-abaxial position and 2 adaxial (on the ventral side). Each vascular bundle is accompanied by some sclerenchymatous elements (Fig. 3, B). These mechanical tissue elements of dill and fennel fruits are more represented in the lateral wings than the rest, while the fruit of cumin sclerenchymatous elements are uniformly arranged right to the vascular bundles. The exception is the carrot fruit whose sclerenchymatous elements reinforce the lateral side thorns. These mechanical elements appear sporadically in anise fruit, discontinuous being weak compared to the rest of the species (Fig. 4, B). Unlike other fruit species, remarkable is, in the carrot fruit, the presence of thorns placed right to the secretive ducts (including the lateral wings) the mesocarp cells taking part in their formation. The vascular bundles are placed right to the protective short unicellular hairs (Bavaru & Bercu, 2002) with a sharp point, arising from a massive epidermal socle slightly sclerified at the base. Anise fruit has several histoanatomical features that distinguish it from other four studied species. Thus each of kidney-shaped mericarps has five slashed ribs and less obvious valecule. The mesocarp is composed of fewer cell layers. The secretive ducts are small, highly elongated and numerous (30-31 per mericarp), some of them located just below the vascular bundles. The epicarp possesses numerous one-celled simple protective hairs, shorter than that of carrots, but in a large number, willing the mericarps almost all around. Endocarp in all species is composed of tangentially elongated cells with straight thicken walls, anise less to that of the radial walls of cells are slightly curled. The seed shows, on the outside, a one-layered tegument with many flattened cells, closely united among themselves and with the endocarp. Anise semen tegument is much more difficult to reveal almost connate with the pericarp. The endosperm consists of large parenchymatous cells (Raven et al., 1992), rich in aleuronal crystals and lipid droplets in all species seeds occupying almost the entire mericarp. The endosperm is well developed in anise fruit, occupying almost the entire mericarp. Compared with the thickness of pericarp, endosperm is most developed in anise fruit, histological being similar to that of other species studied.
Conclusions The 5 considered species, fruits, in histoanatomic terms, follow the same structural plan of organization. In all studied species epicarp and endocarp are similar structured, except for radial walls slightly wavy to fennels. Even if the mesocarp is many-layered in all species, differences arise in terms of number of layers and size of cells that compose it. It is well developed in dill, cumin, carrot and fennel and reduced to anise. The fruits of all 5 studied species possess pore developed vascular bundles in the ribs being more developed to cumin and fennel. Secretive ducts are similar in all studied species differences arise in particular as regards their size (the larger in cumin and the smaller in anise fruit) and their number (30-31 in anise fruit and 6 the other species). The mechanical tissue is represnted by few sclerenchymatous elements, accompanying the vascular bundles. In some of trapezoidal coasted-shaped fruits such as dill and fennel it is present in the lateral wings as well. Annals of RSCB Vol. XVII, Issue 1/2012
270 Epidermal formations meet only the fruit of carrots, represented by emerging (thorns) and simple unicellular protective hairs, the latter present in the fruit of anise as well, being shorter comparatively those of carrot. All studied species are considered food value but are used for therapeutic purposes as well.
References Albert-Puleo, M.: Fennel and anise as estrogenic agents. J. Ethnopharmacol. 2 (4): 33744, 1980..Andrei, M.; Predan, G.M.I.: Practicum de morfologia i anatomia plantelor. Edited by Editura Stiinelor Agricole, Bucureti 2001. Batanouny, K.H.: Anatomy of Plants, 1992. Edited by University Press of Cairo, Cairo. Bavaru, A.; Bercu, R.: Morfologia i anatomia plantelor, 2002. Edited by Ex Ponto, Constana. Bercu, R.: Jianu, D.L.: Practicum de Morfologia i anatomia plantelor, 2003. Edited by Ovidius University Press, Constana. French, D. H.: Ethnobotany of the Umbelliferae. in Heywood, V. H. [ed.], The biology and chemistry of the Umbelliferae, 385-412. Academic Press, London, 1971. Raven, P.H.; Evert, F.R.; Eichhorn, E.S.: Biology of Plants. 5th ed. Edited by Worth Publ. Inc. NY 1992. She, M.; Pu, F.; Pan, Z.; Watson, M.; Cannon, J.F.M.; Holmes-Smith, I.; Kljuykov, E.V.; Phillippe, L.R.; Pimenov, M.G.: Apiaceae. In Flora of China 14: 1205, 2005. Stern, K.R.: Introductory, Plant Biology, sixth edition, 1994. Edited by Wm. C. Brown Communications, Inc., Dubuque (USA). Stevens, P.F.: Angiosperm Phylogeny Website. Version 9, June 2008. http://www.mobot.org/MOBOT/research/AP web/ Watson, L., Dallwitz, M.J.: The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 4th March 2011. http://delta- intkey.com. http://www.complete-herbal.com http://en.wikipedia.org/wiki/Caraway http://en.wikipedia.org/wiki/Daucus_carota http://en.wikipedia.org/wiki/Fennel http://botanical.com