21) Morphology and Anatomy

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Biologia, Bratislava, 62/1: 2—5, 2007

Section Botany
DOI: 10.2478/s11756-007-0006-7
Review

Morphology and anatomy of Echinacea purpurea, E. angustifolia,


E. pallida and Parthenium integrifolium

Ingrid Mistríková & Štefánia Vaverková


Department of Pharmacognosy and Botany, Faculty of Pharmacy, Comenius University, Odbojárov 10, SK–83232
Bratislava 3, Slovakia

Abstract: The Echinacea species are native to the Atlantic drainage area of the United States of America and Canada.
They have been introduced as cultivated medicinal plants in Europe. Echinacea purpurea, E. angustifolia and E. pallida are
the species most often used medicinally due to their immune-stimulating properties. This review is focused on morphological
and anatomical characteristics of E. purpurea, E. angustifolia, E. pallida, because various species are often misidentified
and specimens are often confused in the medicinal plant market.
Key words: anatomy; morphology; medicinal plants; underground parts

Introduction Sloley et al. 2001, Perry et al. 2001; Speroni et al. 2002)
and the chemical profile, in some cases, has been corre-
The Echinacea species are native to North America and lated with a different pharmacological activity (Speroni
are distributed throughout the eastern and central U.S. et al. 2002) enabling more differentiated and appropri-
and southern Canada. Echinacea purpurea, E. angus- ate therapeutic use of these plant species.
tifolia and E. pallida are the species most often used In the products prepared from E. pallida and E.
medicinally. The most widely accepted taxonomic in- angustifolia, the roots are typically used, whereas for E.
terpretation of the genus is that of McGregor (1968), purpurea, juice from the fresh leaves, stems and flowers
in accord with which there are 9 species and 2 varieties. are most often used, though roots are sometimes in-
Formerly considered as a variety, Echinacea pallida, E. cluded (Schulz et al. 2000). Certainly, milled materials
angustifolia and E. purpurea are treated by McGregor from different Echinacea species are indistinguishable
as separate species on the basis of morphological cri- by morphological characteristics.
teria, cultivation and hybridization experiments, and Since commercial roots of E. purpurea have been
cytological features (Foster 1991). Various species are adulterated with Parthenium integrifolium, and E. an-
often misidentified. There are many morphological sim- gustifolia roots have frequently been mixed with E. pal-
ilarities between the species, especially between E. an- lida, the study of morphological and anatomical char-
gustifolia and E. pallida, and it has been shown that acters of Echinacea roots has become of great impor-
specimens are often confused in the medicinal plant tance. Important morphological and anatomical char-
market. E. angustifolia was classified as a variety of acteristics of Echinacea plants and Parthenium integri-
E. pallida (Bradley 1992). Many of the early studies folium are described for the purpose of pharmacognos-
on E. angustifolia and E. purpurea were probably ac- tical identification of the drugs.
tually conducted on E. pallida, and studies published
before 1987 must be viewed with caution, consider- Morphological description
ing the actual species evaluation (Bauer et al. 1987;
Schumacher & Friedberg, 1991). The Echinacea prod- The word “echinacea” comes from the Greek word
ucts have also been frequently adulterated with another “echinos”, which means sea urchin or hedgehog. It has
plant of similar appearance Parthenium integrifolium, probably got this name from the prickly spikes found
which has none of the pharmacological effects (Hobbs on the flower head.
1989; Bradley 1992). This has resulted in confusion over Echinacea species are herbaceous perennials of the
the therapeutic properties of each species (Bauer et al. Asteraceae/Compositae family reaching 10–60 cm of
1987). height. The stem ascends either from a vertical taproot
Nevertheless, significant differences, both qualita- (E. angustifolia) or branched, fibrous roots (E. pur-
tive and quantitative, in the chemical composition of purea). Echinacea may have either simple or branched
the three species were reported (Mazza & Cotrell 1999; stems. As members of the Asteraceae family, each


c 2007 Institute of Botany, Slovak Academy of Sciences
Morphology and anatomy of Echinacea species 3

“flower” or daisy-like head unit is actually a conglomer- 3 cm, long, 0.5–1 cm broad, purplish; pales 0.9–1.3 cm
ation of many tiny florets. The inner (disc) florets end in long, awn half as long as body; disc corollas 0.45–0.55
spines, and are surrounded by droopy outer (ray) florets cm long, lobes 0.1 cm long; achene 0.4–0.45 cm long,
with teeth at their ends. Echinacea genus is character- pappus a low crown of equal teeth; pollen grains yel-
ized by spiny flowering heads, with an elevated recep- low, 19–21 µm in diameter; chromosome number n = 11
tacle which form the “cone”. The colour of ray florets (McGregor 1968).
ranges from white, through pink to deep purple, with
yellow reserved for E. paradoxa var. paradoxa. Disc flo- Parthenium integrifolium L.
rets may be red-brown to green. The leaf shape varies Stems branched, 60–130 cm high; leaves egg-shaped,
from lanceolate to ovate. eliptical, dentate to serrate edge, 10–25 cm long, 6–
Echinacea plants are resilient and drought resis- 12 cm broad, slightly hairy; the number of flowers in
tant, but they grow slowly (Awang & Kindack 1991; the panicle varies; rays spreading 0.15–0.3 cm long, 0.1
Houghton 1994). cm wide; disc corollas 0.1–0.15 cm long, sterile; pollen
grains white, 12–22 µm in diameter; chromosome num-
Echinacea angustifolia DC. var. angustifolia ber n = 9.
Stems simple or occasionally branched, 10–50 cm high, In contrast to Echinacea species the taproot of
smooth or hirsute below, hirsute or tuberculate-hispid Parthenium integrifolium is short, only 3–8 cm long, to
above; leaves oblong-lanceolate to elliptical, entire, dark 4 cm in diam, bulbform, thickened, black. The branches
green, tuberculate hirsute to tuberculate-hispid; basal arise from the bottom part of the taproot. The lateral
leaves short to long-petiolate, 5–27 cm long, 1–4 cm roots are partially shiny.
broad, lower cauline leaves petiolate, 4–15 cm long,
0.5–3.8 cm broad, upper cauline leaves sessile, acute; Anatomical description
heads 1.5 cm high, 1.5–2.5 cm broad exclusive of ligules;
phyllaries in 3–4 series, lanceolate, acute, entire, 0.6– The anatomical descriptions of the underground parts
1.1 cm long 0.2–0.3 cm wide, tuberculate-hirsute or of Echinacea were published in the late eighties of the
tuberculate-hispid; rays spreading, 2–3.8 cm long, 0.5– last century (Bauer & Wagner 1987; Heubl et al. 1988).
0.8 cm wide, white, pinkish or purplish; disc corollas The increasing interest in Echinacea and the adulter-
0.6–0.85 cm long, lobes 0.12–0.2 cm long; achenes 0.4– ations that had been observed inspired Bauer et al.
0.5 cm long, pappus a toothed crown; pollen grains yel- (1987) and Heubl et al. (1988) to examine the roots of
low, 19–26 µm in diameter; chromosome number n = 11 E. purpurea, E. angustifolia, E. pallida and of Parthe-
(McGregor 1968). nium integrifolium. Their key for the identification of
Echinacea-powders and the detection of Parthenium
Echinacea pallida (Nutt.) Nutt. became standard (Bauer & Wagner 1990).
Stems simple, rarely branched, 40–90 cm high, sparsely Heubl et al. (1988) showed that for a precise iden-
hirsute below, more densely so above; leaves oblong- tification of the roots besides the differentiation of the
lanceolate to long-elliptical, entire, dark green, hirsute exodermis cells, especially the sclerenchymatous struc-
on both surfaces, triple-neined; basal leaves 10–35 cm tures (stone cells, sclerotic fibres) and the occurrence
long, 1–2.5 cm broad, acute, petiolate below to sessile and distribution of phytomelan are of great importance.
above; phyllaries lanceolate to narrowly oblong, 0.8–1.7 In addition, specific arrangement of the vessels in the
cm long 0.2–0.4 cm broad, hirsute, ciliate, 3–4 series, cross – and longitudinal sections can be used for the
gradually passing into the echinaceous pales; rays re- characterisation of the species.
flexed, 4–9 cm long, 0.5–0.8 cm broad, purplish, pink, Länger (2001) considered the application of the key
or white; pales 1–1.3 cm long, body 0.8–1 cm long, awn of Heubl failing, mainly due to confusing terminology,
0.25–0.34 cm long; disk corollas 0.8–1 cm long, lobes lack of detailed drawings, and some of the proposed
0.2–0.3 cm long; achenes 0.37–0.5 cm long, glabrous, differencing characters (size and shape of the exoder-
pappus a toothed crown, teeth about even, longest 0.1 mal cells and the different localization of the secretory
cm; pollen grains white, 24–28 µm in diameter; chro- cavities). The author supposes that the anatomical dif-
mosome number n = 22 (McGregor 1968). ferences between the roots of E. angustifolia, E. pallida
and E. atrorubens are not sufficient for differentiation,
Echinacea purpurea (L.) Moench however, root and rhizome of E. purpurea and the root
Stems erect, stout, branched, hirsute or glabrous, 60– of Parthenium integrifolium appear well characterized.
180 cm high; basal leaves ovate to ovate-lanceolate, Radix Echinaceae consists of the fresh or dried
acute, coarsely or sharply serrate, petioles up to 25 roots of Echinacea angustifolia D.C. var. angustifolia
cm long, blades to 20 cm long and 15 cm wide, blade or its variety strigosa McGregor, or E. pallida (Nutt.)
abruptly narrowing to base, often cordate, decurrent on Nutt. (Asteraceae).
petiole, 3–5 veined; cauline leaves petiolate below, ses- The Länger’s publication is the last one dealing
sile above, 7–20 cm long, 1.5–8 cm broad, coarsely ser- with the anatomy of the underground parts of Echi-
rate to entire, rough to the touch on both surfaces; phyl- nacea species. A description of the microscopic char-
laries linear-lanceolate, attenuate, entire, pubescent on acteristics of a cross-section of the aerial parts of the
outer surface, ciliate, passing into the chaff; heads 1.5– Echinacea plants is not available.
4 I. Mistríková & Š. Vaverková

Anatomy of the underground parts of the exodermis cells, sclereids and secretory cavities
is in average bigger in E. pallida.
Echinacea angustifolia Sclereids coated with phytomelanin are solitary or
Root: The transverse section shows a thin outer bark in groups. Solitary sclereids are more or less isodia-
separated from a wide xylem by a distinct cambium metric with a diameter up to 80 µm, whereas those
line. Brown pigmented exodermis or, in older roots in groups are slender and elongated with a diameter up
a brown cork, represents the outermost layer of the to 50 µm and the length of approximately 500 µm. The
root. Cork is composed of several rows of thin-walled lignified fibres are present only in the periphery of the
cells containing yellowish brown pigment. Cortex is cortex and phytomelanin is often absent.
parenchymatous. The secretory cavities, often in cross section tan-
Secretory cavities, 80–150 µm in diameter and con- gentially elongated, reach dimensions up to 600 µm.
taining yellowish orange oleorisin are present only out- The structure of the xylem fibres is like in E. an-
side the central cylinder, in the cortical parenchyma. gustifolia (Länger 2001).
Sclereids with phytomelanin are found alone or in Powder: Frequently sclereids with phytomelanin, mostly
groups of two or three, or sometimes up to ten, varying in elongated and multiseriate groups; only few soli-
considerably in size and shape from rounded to rect- tary sclereids; few bundles of fibres without phytome-
angular to elongated and fibre-like, up to 50 µm in di- lanin; fragments of exodermis; colourless parenchyma;
ameter and up to 300 µm in length, with intercellular tracheas with reticulate thickening or bordered pits
spaces. Phytomelanin fills the triangular intercellular (Länger 2001).
spaces around the sclereids, making them star-shaped
in cross section. In longitudinal view, these sclereids are Echinacea purpurea
characterized by numerous pit channels and a rather Root: The root is covered with dark brown exodermis
small lumen. consisting of polygonal cells. The cortical parenchyma
The secondary phloem and the secondary xylem has thinner walls than those found in the secondary
are composed of very narrow strands of vascular tis- phloem.
sue separated by wide, non- lignified medullary rays. Secretory cavities, filled with orange to brown mat-
Within the radial rows the vessels are arranged in ter are small, 50–120 µm long. They are located at the
small groups separated by parenchyma. The vessels border between cortex and secondary phloem, mostly
are 25–75 µm in diameter and have secondary walls in groups of 3 or 4 at radial position of the groups of
with reticulate or scalariform thickenings but occa- tracheas within the xylem part. The total number of
sionally with spiral or annular thickenings or bordered secretory canals varies from 12 to 20.
pits. Frequently fibres, usually without phytomelanin, Stone cells occurring rarely in the cortex parenchyma,
are associated with the vessel members. In longitudi- are mostly single and phytomelanin is often absent
nal section, a thinner cell wall and slender shape with (Heubl 1988). According to Länger (2001), sclereids and
pointed ends distinguishes these fibres from sclereids. phytomelanin are absent in the root.
Within the secondary xylem, sclereids are located in The vessels and fibres of the secondary xylem are
the medullary rays only, whereas secretory cavities are grouped into cuneiform bundles tapering towards the
scattered throughout the parenchyma tissue. Surround- centre of the root. In younger roots more parenchyma
ing the small pith, sparse groups of primary xylem is located within these areas, whereas in older ones the
with narrow vessels are situated at the inner end of secondary xylem becomes a completely solid structure.
the medullary rays (Heubl et al 1988; Länger 2001). Small groups of primary xylem are located at the inner
Rhizome: Apart from the different arrangement of the end of the medullary rays, outside of the central pith. In
primary and secondary elements, cell structures are the longitudinal section, pitted and scalariform vessels
identical with those of the root (Länger 2001). are found associated with elongated and pointed fibres.
Powder: Powdered rhizome and roots are brown with Rhizome: The pigmented exodermis consists of polygo-
a slight aromatic odour and initially a sweet taste, nal cells. The cortex is characterized by slightly thick-
quickly becoming bitter and leaving a tingling sensa- ened parenchyma cells with small triangular intercellu-
tion on the tongue. Thin-walled polygonal cork cells lar spaces.
with red-brown contents; lignified reticulately thick- Sclereids with a diameter of approximately 60 µm
ened vessels. Frequently sclereids with phytomelanin, and a length up to 300 µm occur singly or in small
mostly in elongated multiseriate groups; occasional groups. Phytomelanin usually covers the cell wall of
bundles without phytomelanin. Fragments of secretory these sclereids and fills in the triangular intercellular
cavities with reddish brown contents; abundant thin- spaces around them, making their star-shaped appear-
walled parenchyma with spherocrystalline masses of in- ance in cross section.
ulin (Länger 2001). Groups of fibres are placed like a cap outside of
the phloem areas. Within the cambial line the vascu-
Echinacea pallida lar bundles are separated and arranged circumferen-
Rhizome and root: The main structure and the cellular tially. In the secondary xylem, the parenchyma between
elements are very similar to E. angustifolia, but, pos- and within the cambial bundles is replaced by fibres
sibly due to the doubled chromosome number, the size and thickened isodiametric cells, creating a solid ring
Morphology and anatomy of Echinacea species 5

of xylem tissue without medullary rays. Most vessel References


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of approximately 40–60 µm occur in the cortical McGregor R. 1968. The taxonomy of the genus Echinacea (Com-
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fibres, most sclereids are yellow. The secondary xylem dardization: analytical methods for phenolic compounds and
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Arzneimmittel Forschung 41: 141–147.
medullary rays contain very large, often ruptured cells. Sloley B.D., Urichuk L.J., Tywin C., Pang P.K. & Shan J.J. 2001.
Within the secondary xylem, sclereids only occur in the Comparison of chemical components and antioxidants capac-
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Compared to the roots of Echinacea spp., the root 849–857.
Speroni E., Govoni P., Guizzardi S., Renzulli C. & Guerra M.C.
of Parthenium integrifolium has the smallest amount 2002. Anti-inflammatory and cicatrizing activity of Echinacea
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row fibres free of phytomelanin deposits may be found
within the region of the primary xylem (Länger 2001). Received Nov. 23, 2005
Accepted May 4, 2006
Powder: Dark brown pigmented exodermis with rect-
angular cells. Numerous sclereids with phytomelanin-
coating in groups from 5 to 30 cells. Multiseriate frag-
ments of phytomelanin-coated fibres with associated
pitted vessels. Colourless parenchyma; infrequent bun-
dles of fibres.

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