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T. K.


Edible Medicinal
and Non-Medicinal
Volume 11,
Modified Stems, Roots, Bulbs
Edible Medicinal and Non-Medicinal
T.K. Lim

Edible Medicinal and

Non-Medicinal Plants
Volume 11, Modified Stems, Roots,
ISBN 978-3-319-26061-7 ISBN 978-3-319-26062-4 (eBook)
DOI 10.1007/978-3-319-26062-4

Library of Congress Control Number: 2011932982

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Special thanks for the use of digital images are accorded to Cecilia Lafosse
(CIP) and Ezeta Fernando (ex CIP), International Potato Centre (CIP) for oca
(Oxalis tuberosa). Frank Van Keirsbilck for mauka (Mirabilis expansa) and
GF Chung for Indian goosegrass (Eleusine indica).


The author and publisher of this work have checked with sources believed to
be reliable in their efforts to confirm the accuracy and completeness of the
information presented herein and that the information is in accordance with
the standard practices accepted at the time of publication. However, neither
the author nor publishers warrant that information is in every aspect accurate
and complete and they are not responsible for errors or omissions or for con-
sequences from the application of the information in this work. This book is
a work of reference and is not intended to supply nutritive or medical advice
to any individual. The information contained in the notes on edibility, uses,
nutritive values, medicinal attributes and medicinal uses and suchlike included
here are recorded information and do not constitute recommendations. No
responsibility will be taken for readers’ own actions.


Introduction ........................................................................................... 1


Iris x germanica ..................................................................................... 27


Stachys affinis ........................................................................................ 41


Maranta arundinacea ............................................................................ 47


Nelumbo nucifera .................................................................................. 55


Mirabilis expansa ................................................................................... 110


Nymphaea odorata ................................................................................. 114

Nymphaea x rubra ................................................................................. 118
Victoria amazonica................................................................................. 123


Diuris semilunulata................................................................................ 128

Diuris sulphurea .................................................................................... 130
Pterostylis curta ...................................................................................... 133
Pterostylis pedunculata .......................................................................... 136

x Contents


Oxalis tuberosa....................................................................................... 139


Piper methysticum .................................................................................. 147


Chrysopogon zizanioides........................................................................ 197

Eleusine indica ....................................................................................... 228


Morinda officinalis ................................................................................ 237


Eurycoma longifolia .............................................................................. 250

Medical Glossary................................................................................... 277

Scientific Glossary ................................................................................. 354

Common Name Index ........................................................................... 378

Scientific Name Index ........................................................................... 385


This book continues as volume eleven of a multi- edible fruits or flowers have been covered in ear-
compendium on Edible Medicinal and Non- lier volumes and for those better known for other
Medicinal Plants. It covers plants with edible non-reproductive plant parts will be covered in
modified storage subterranean stems (corms, rhi- latter volumes.
zomes, stem tubers) and unmodifed subterranean As in the preceding ten volumes, topics cov-
stem stolons, above-ground swollen stems and ered include: taxonomy (botanical name and syn-
hypocotyls, storage roots (tap root, lateral roots, onyms); common English and vernacular names;
root tubers) and bulbs that are eaten as conven- origin and distribution; agro-ecological require-
tional or functional food as vegetables and spices, ments; edible plant part and uses; plant botany;
as herbal teas, and may provide a source of food nutritive and medicinal/pharmacological proper-
additive or neutraceuticals. A list of such edible ties with up-to-date research findings;traditional
plant species from families Acanthaceae to medicinal uses; other non-edible uses; and
Zygophyllaceae are presented in a tabular form selected/cited references for further reading.
and 32 such edible species from the families A corm or bulbotuber is defined as a short,
Alismataceae, Amaryllidaceae, Apiaceae, vertical, swollen underground plant stem that
Araceae, Araliaceae, Asparagaceae, Asteraceae, serves as a storage organ used by some plants to
Basellaceae, Brassicaceae and Campanulaceae survive unfavourable adverse periods. It bears
had been covered in detail in preceding volume 9. membranous or scaly leaves and buds. Some
Nineteen edible species from the families examples of plants with edible corms are found
Amaranthaceae, Cannaceae, Cibotiaceae, in Amorphophallus spp., Colocasia esculenta
Convolvulaceae, Cyperaceae, Dioscoreaceae, (taro), Eleocharis dulcis (Chinese water chest-
Euphorbiaceae and Fabaceae had been covered in nut), Sagittaria spp. (arrowhead or wapato) and
detail in volume 10. This volume 11 covers in Xanthosoma spp. (cocoyam or tannia). Corms
detail 18 edible species in the families Iridaceae often give rise to many small secondary corms or
(1), Lamiaceae (1), Marantaceae (1), cormlet called cormels at the end of very short
Nelumbonaceae (1), Nyctaginaceae (1), stolons.
Nymphaeaceae (3), Orchidaceae (4), Oxalidaceae Rhizome is a modified subterranean stem of a
(1), Piperaceae (1), Poaceae (2), Rubiaceae (1) plant that is usually found underground, produc-
and Simaroubaceae (1). Other species from these ing roots and shoots. It is used by the plant as
families with edible modified stems, roots and storage organ and whole rhizome or pieces of the
bulbs are listed in Table 1. Many plants with such rhizome serves as vegetative propagules to give
edible plant parts that are better known for their rise to new plants. Examples of plants with edible

© Springer International Publishing Switzerland 2016 1

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_1
2 Introduction

rhizomes include gingers (Zingiber spp.), tur- leaves or leaf bases that serve as food storage
meric (Cucurma longa), greater galangal (Alpinia organs during dormancy and enable the plant to
galanga), lesser galangal Alpinia officinarum), survive through adverse periods. The fleshy
sand ginger or kencur (Kampferia galanga), lotus leaves are arranged in a concentric manner. Bulbs
root (Nelumbo nucifera), Typha spp., fingerroot can be tunicate i.e. with membranous papery cov-
(Boesenbergia rotunda) and arrowroot (Maranta ering (scale leaves) or tunic that protects the inner
arundinacea). fleshy scale leaves from drying and mechanical
A stem tuber is a modified plant storage organ injury. Examples of tunicate bulbs are the
that is formed from thickened rhizome or stolon. Alliums, onions, leeks, hyacinth and tulips. In
The tops or sides of the tuber produce shoots that imbricate or non-tunicate bulbs, the fleshy scale
grow into typical stems and leaves and the under- leaves are not in concentric rings but are loosely
sides produce roots. The stem tuber has all the arranged or spreading, overlapping one another
parts of a normal stem, including nodes (eyes) and at the margin. Such a bulb is not a compact body
internodes. A stem tuber may start off as an and not usually covered by a common tunic.
enlargement of the hypocotyls of the seedling and Examples are the garlic (Allium sativum) and
may include the epicotyl or upper section of the some Lilium lilies.
root as is in the case of maca (Lepidium meyenni). Tap root is the true main root of the plant and in
More commonly as in Plectranthus esculenta in some species the tap root is modified and fleshy,
the Lamiaceae family, numerous tubers are rich in stored nutrients; they may or may not be
formed on short stolons that arise from the base of fused with the hypocotyl or basal stem tissues and
the stem, or as in potatoes tubers are formed as maybe napiform, globose, conical, fusiform or
enlarged stolons thickened and enlarged into stor- cylindrical in shape. Notable examples of plants
age organs. In some Cyperus species e.g. tigernut with edible tap roots are Abelmochus spp., beet
or chufa (C. esculentus), the stolons end with the (Beta vulgaris), rutabaga, turnip, Bunium persi-
growth of tubers that can give rise to new plants. cum, burdock, carrot, radish and daikon, celeriac,
Other striking examples of plants with stem tubers jicama and ahipa (Pachyrhizus spp.), parsnips,
include hog potato or groundnut (Apios ameri- parsley, skirret (Sium sisarum), bush potato (Vigna
cana), Jerusalem artichoke or sunchoke lanceolata), salsify (Tragopogon porrifolius),
(Helianthus tuberosus), earthnut pea (Lathyrus black salsify (Scorzonera hispanica), tongkat Ali
tuberosus), oca or New Zealand yam (Oxalis (Eurycoma longifolia) and many others. Plants
tuberosa), Chinese artichoke or crosne (Stachys with edible root tubers or tuberous roots with
affinis), mashua or añu (Tropaeolum tuberosum) enlarged root and lateral roots function as storage
and ulluco (Ullucus tuberosus). In Botany, a sto- organs, lacking nodes, internodes and adventious
lon is an horizontal modified stem arising from buds. Notable examples include pignut or earthnut
the base of a plant that produces new plants from (Conopodium majus), sweet potato (Ipomoea bata-
buds at its tip or nodes and forms adventitious tas), desert yam (Ipomoea costata), cassava or yuca
roots at the nodes, it can be creeping above the or manioc (Manihot esculenta), yams (Dioscorea
ground surface or underground. An example of a spp.), mauka or chago (Mirabilis expansa), bread-
plant with edible stolon is Imperata cylindrica. root, tipsin, or prairie turnip (Psoralea esculenta)
However, some botanists used the term stolons for and yacón (Smallanthus sonchifolius).
stem branches that arise from the base of the stem Most terrestrial orchids are rhizomatous or
that creeps above the ground and those that creeps have corms or tubers. Many orchid species pro-
horizontally underground as rhizomes. An exam- duce edible tubers and roots while comparatively
ple of a plant with swollen, above-ground storage fewer species (e.g. Cymbidium canaliculatum
stem is the kohlrabi. and Dendrobium tarberi) produce edible pseudo-
Bulb is a much reduced underground stem bulbs. These tubers contain a nutritious, starchy
bearing at its apex a growing or floral primor- polysaccharide called glucomannan. These
dium surrounded by thick, fleshy modified scale tubers provide a starchy flour called salep which
Introduction 3

Table 1 Plants with edible modified stems, roots and bulbs in the families: Iridaceae, Lamiaceae, Marantaceae,
Nelumbonaceae, Nyctaginaceae, Nymphaeaceae, Orchidaceae, Oxalidaceae, Piperaceae, Poaceae, Rubiaceae and
Family Scientific Name Names Edible Part Use Reference
Iridaceae Crocus sativus L. Saffron Roots are eaten roasted Hedrick (1972),
Kunkel (1984),
Morton (1976),
Facciola (1990),
and Lim (2014)
Iridaceae Iris cristata Aiton Dwarf Crested Iris Roots used as a spice. Hedrick (1972),
Frequently chewed by Tanaka (1976),
local people to alleviate Kunkel (1984),
thirst. When first and Coffey (1994)
chewed, the roots have
a pleasant sweet taste,
within a few minutes
this changes to a
burning sensation far
more pungent than
Iridaceae Iris x germanica L. Bearded Iris, Flag, Orris Root used as spice, Chase (1900),
Root, German Iris, flavouring in ice cream, Parmentier (1781)
Florentine Iris confectionery and cited by
baked goods; starch Freedman (2009),
used as bread flour Uphof (1968),
Morton (1976),
Facciola (1990),
Bender (2009),
and Surhone et al.
Iridaceae Iris pallida Lam. Sweet Iris, Dalmation Orris oil from rhizome Morton (1976)
Iris used to flavour soft
drinks, candy and
chewing gum
Iridaceae Iris setosa Pall. ex Link Hiogi-Ayame Rhizomes eaten or used Uphof (1968),
as source of starch Hedrick (1972),
Tanaka (1976),
and Facciola
Iridaceae Morea fugax Uintjie Bulbous root eaten Hedrick (1972),
(D. Delaroche) Jacq. roasted, boiled or Fox et al. (1982),
stewed with milk and Facciola
Iridaceae Romulea bulbocodium NF Bulbous root eaten Fairchild (1930)
(L.) Sebast. & Mauri
Iridaceae Tigrida pavonia (L. f.) Common Tiger Flower, Roasted starchy corms Uphof (1968) and
Redouté Jockey Cap, Mexican used as food by Facciola (1990)
Shellflower, Peacock Mazatecs and other
Flower Indian tribes in Mexico
Lamiaceae Callicarpa rubella Gopura Esing (Mishing) Bark and roots chewed Patiri and Borah
Lindl. Bonmala (Assamese) like betel nut; roots (2007) and
eaten in Meghalaya Sawian et al.
4 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Lamiaceae Callicarpa vestita Wall. Yarpo Esing (Mishing) Bark and roots chewed Patiri and Borah
ex C.B.Clarke like betel nut (2007)
Lamiaceae Clerodendrum fragrans Fragrant Glorybower; Roots dried, cooked Hu (2005)
(Vent.) R.Br. = Chou Mo Li, Chou Mu with pork to strengthen
Clerodendrum chinense Dan (Chinese) elderly people and to
(Osbeck) Mabb, remove pain and
stiffness of muscles and
Lamiaceae Clerodendrum serratum Phelang Riho Roots eaten in Karbi, Kar and
(L.) Moon = Rotheca (Assamese) Assam Borthakur (2008)
serrata (L.) Steane &
Lamiaceae Coleus blumei Benth. = Coleus, Painted Nettle, Tubers eaten Burkill (1966)
Plectranthus Sayabana, Jacob’s Coat and Facciola
scutellarioides (L.) (1990)
Lamiaceae Coleus dazo A. Chev. = Daju, Rizuka Starchy root peeled, Tanaka (1976)
Plectranthus esculentus boiled, served and and Facciola
N.E.Br. eaten or pickled (1990)
Lamiaceae Coleus parviflorus African Potato, Country Tubers eaten like Tanaka (1976),
Benth. = Plectranthus Potato potatoes Ochse and van
rotundifolius (Poir.) den Brink (1980)
Spreng. and Facciola
Lamiaceae Coleus tuberosus African Potato, Country Tubers usually eaten Ochse and van
(Blume) Benth. = Potato steamed or cooked with den Brink (1980)
Plectranthus rice In Indonesia
rotundifolius (Poir.)
Lamiaceae Eriophyton wallichii Mian Shen (Chinese) Roots used in food in Hu (2005)
Benth. ex Wall. north-western Yunnan
Lamiaceae Leonurus sibiricus L. Siberian Motherwort In China, the roots are Burkill (1966),
cooked with pork Altschul (1973),
Tanaka (1976),
and Facciola
Lamiaceae Lycopus europaeus L. Gypsywort, Water China: root eaten. Read (1946) and
Horehound Manchuria: starchy Baranov (1967)
tubers eaten
Lamiaceae Lycopus lucidus Turcz. Lycopos, Bugleweed; Di Underground rhizomes Hu (2005)
ex benth. Gua Er Miao (Chinese) eaten in northern china
and Yunnan
Lamiaceae Lycopus uniflorus Northern Bugleweed White tubers eaten raw Fernald et al.
Michx. in salads, boiled, (1985) and
pickled or added to Facciola (1990)
soups and stews
Lamiaceae Phlomis tuberosa L. Tuberous Jerusalem Roots eaten by the Hedrick (1972)
Sage; Bodmon Sok Kalmucks in Eurasia and Facciola
Lamiaceae Plectranthus barbatus Indian Coleus Tubers eaten Jansen (1996)
Introduction 5

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Lamiaceae Plectranthus edulis Galla Potato Tubers eaten Jansen (1996)
Lamiaceae Plectranthus esculentus Livinstone Potato, Kaffir Stem tubers eaten, Jansen (1996),
N.E. Br. Potato eaten raw or boiled and Phillips and Rix
eaten as vegetables (1993), van Wyk
(2006), and
Codex (2014)
Lamiaceae Plectranthus Madagascar Spur Flower Tubers eaten in Tanaka (1976)
madagascariensis Madagascar and Facciola
(Pers.) Benth. (1990)
Lamiaceae Plectranthus Chinese Potato, Coleus Young aromatic tubers Jansen (1996) and
rotundifolius (Poir.) Potato, Hausa Potato used in soup and Codex (2014)
Spreng vegetable dishes
Lamiaceae Solenostemon Huasa Potato, Fra-Fra Nigeria (Kano State, Dalziel (1955),
rotundifolia (Poir.) Potato; Tumuku, Tamaka northern): tuber eaten Fox et al. (1982),
J.K. Morton. = (Hausa) like potato Mortimore
Plectranthus (1989), Facciola
rotundifolius (Poir.) (1990), and
Spreng Codex (2014)
Lamiaceae Stachys adulterina Hubei Artichoke; Di Can Root tubers used as Hu (2005)
Hemsl. Zi (Chinese) vegetables, cooked or
Lamiaceae Stachys affinis Bunge Chinese Artichoke; Cao As above Phillips and Rix
Shi Can (Chinese) (1993) and Hu
(2005), Codex
Lamiaceae Stachys chinensis Bunge Hyssopleaf Hedgenettle Manchuria: rhizome Baranov (1967)
ex Benth eaten
Lamiaceae Stachys sieboldi Miq. Crosnes, Chinese Japan: tubers salted or Read (1946),
Artichoke, Japanese preserved in plum Facciola (1990),
Artichoke vinegar Van den Bergh
(1996), and
Codex (2014)
Marantaceae Calathea allouia Guinea Arrowroot, Root tubers boiled and Facciola (1990),
(Aublet) Lindl. Leren, Sweet Corn Tuber eaten like potato Groen et al.
(1996), and
Codex (2014)
Marantaceae Halopegia blumei Jelantir (Javanese), Patat Tubers eaten cooked or Groen et al.
(Körn) K Schum. (Sundanese) Dong Nam roasted (1996), Ochse
(Vietnam) and van den Brink
(1980), and
Ochse and van
den Brink (1980)
Marantaceae Maranta arundinacea L. Arrowrot, Tora Alu, Tha Rhizomes are source of Ochse and van
Lairusa, Hnathel, arrowroot, eaten den Brink (1980),
Hpogimbai (Assamese) cooked or raw Tuber Facciola (1990),
Khaita Alu (Boro) eaten both raw and Villamayor Jr and
Nginti Ali (Mishing) boiled, starch from Jukema (1996),
rhizome Hu (2005), Patiri
and Borah (2007),
Medhi and
Borthakur (2012),
and Codex (2014)
6 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Marantaceae Maranta dichotoma Mohtra Reed, Sitalpati Tuber eaten both raw Medhi and
(Roxb.) Wall. = Plant; Tha Lairu, and boiled Borthakur (2012)
Schumannianthus Hnathel, Hpogimbai
dichotomus (Roxb.) (Assamese)
Marantaceae Phrynium capitatum Packing Leaf Root tuber eaten in Sawian et al.
Willd. = Phrynium Meghalaya (2007)
pubinerve Blume
Marantaceae Thalia geniculata L. Swamp Lily Rhizomes baked and Tanaka (1976)
eaten or made into a and Facciola
kind of arrowroot (1990)
Nelumbonaceae Nelumbium speciosum Pink Water Lily India: root and seeds Gammie (1902)
Willd. eaten and Watt (1908)
Nelumbonaceae Nelumbo lutea Pers. American Lotus, Water Large tubers, when Facciola (1990)
Chinquapin baked, are sweet and and Saunders
mealy with a flavour (1920)
somewhat like a
Nelumbonaceae Nelumbo nucifera Lotus, Lotus Root Root eaten raw or Burkill (1966),
Gaertn. cooked, sliced used in Cribb and Cribb
stir-fries, soups, stews (1987), Facciola
or fried as a garnish or (1990), Phillips
side dish. Sliced pieces and Rix (1993),
can be candied or Ong (1996), Hu
pickled. Lotus root (2005), Santich
flour is starch and can et al. (2008), van
be used to make Wyk (2006), and
desserts Codex (2014)
Nyctaginaceae Abronia latifolia Yellow Sand Verbena Root edible Yanovsky (1936)
Eschsch. and Facciola
Nyctaginaceae Boerhavia coccinea Tar Vine, Hog Weed Bland fibrous tap root Cribb and Cribb
Mill. eaten (1987) and
Harden (1990)
Nyctaginaceae Boerhavia diffusa L. Hog Weed, Horse Fleshy portion of thick Hu (2005)
Purslane; Zhu Er Yan, roots roasted and eaten,
Huang Xi Xin (Chinese) sweetish and nutritious
Nyctaginaceae Boerhavia spp. Tar Vines Bland fibrous tap root Low (1991)
Nyctaginaceae Mirabilis expansa (Ruiz Mauka, Chago Tuber dried, boiled or Tanaka (1976),
& Pav.) Standl. fried, eaten as Popenoe et al.
vegetable (1989), Facciola
(1990), Bermejo
and Leon (1994),
and Codex (2014)
Nymphaeaceae Euryale ferox Salisb. Chicken Head, Fos Nut In China, roots and Read (1946) and
seeds eaten Facciola (1990)
Nymphaeaceae Nuphar advena R.Br. Common Spatterdock Rootstock eaten raw, Uphof (1968),
roasted or cooked with Hedrick (1972),
meat and Facciola
Introduction 7

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Nymphaeaceae Nuphar luteum Sibth. & Yellow Water Lily Rootstock boiled as Hedrick (1972),
Sm. vegetable Fernald et al.
(1985), and
Facciola (1990)
Nymphaeaceae Nuphar polysepala Cow Lily, Spatterdock, Rich starchy rhizome Schofield (2003)
Engelm. = Nuphar lutea Pond Collard used as survival food,
subsp. polysepala after boiling, roasting
(Engelm.) E.O. Beal. or baked and skin
Nymphaeaceae Nuphar pumilum Yellow Pond Lily; Ping Young tender rhizomes Hu (2005)
(Timm.) DC. Peng Cao (Chinese) used as potherb in
Yunnan and Hubei
Nymphaeaceae Nymphaea lotus L. Egyptian Lotus, White Tubers edible Tanaka (1976),
Lotus Kunkel (1984),
and Facciola
Nymphaeaceae Nymphaea stellata Blue Lotus Of India Rhizomes eaten raw or Uphof (1968),
Willd. = Nymphaea roasted Hedrick (1972),
nouchali Burm.f. Tanaka (1976),
and Facciola
Nymphaeaceae Nymphaea alba L. White Lotus, European In France, root Parmentier (1781)
White Waterlily recommended as a (cited by
famine food after Freedman (2009))
cooking in water and
being flavoured
Nymphaeaceae Nymphaea caerulea Blue Lotus Of Egypt, Starchy tubers eaten Tanaka (1976),
Savigny = Nymphaea Blue Water Lily boiled or roasted Fox et al. (1982),
nouchali var. caerulea and Facciola
(Savigny) Verdc. (1990)
Nymphaeaceae Nymphaea edulis DC. Red Water Lily; In India (Madras Shortt
Shunguneer Pushpum Presidency), roots and (1887–1888)
(Tamil); Koteka, seeds cooked and eaten
Kalharamu (Telugu)
Nymphaeaceae Nymphaea gigantea Giant Water Lily Tuberous rootstock Cribb and Cribb
Hook. eaten (1987)
Nymphaeaceae Nymphaea lotus var. Red Water Lily Root eaten baked or Paton and Dunlop
pubescens (Willd) boiled with salt added (1904)
Hook.f. &
Thomson = Nymphaea
pubescens Willd.
Nymphaeaceae Nymphaea lotus L. Egyptian Lotus, White In upper Guinea Africa, Gammie (1902),
Lotus; Bado (Hausa); root used as a famine Watt (1908),
Dambi (Kanuri) food, being either Irvine (1952),
roasted in ashes or Uphof (1968),
dried before being Hedrick (1972),
ground into flour. In Tanaka (1976),
Nigeria (Kano State, Pongpangan and
northern), rhizome and Poobrasert
seeds eaten. In India (1985),
(Bombay Presidency), Mortimore
roots and seeds eaten (1989), and
Facciola (1990)
8 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Nymphaeaceae Nymphaea nouchali Boga Bhet, Seluk Rhizomes/roots eaten Van den burgh
Burm.f. (Assamese) raw or cooked as (1994) and Patiri
vegetable in Assam and Borah (2007)
Nymphaeaceae Nymphaea odorata Frgarny Water Lily Tubers edible Tanaka (1976),
Aiton Kunkel (1984),
and Facciola
Nymphaeaceae Nymphaea rubra Roxb. Ronga Bhet, Mokua, As above Patiri and Borah
ex Andrew Seluk (Assamese) (2007)
Nymphaeaceae Nymphaea spp. Waterlilies Fibrous tubers edible Cribb and Cribb
(1987) and Low
Nymphaeaceae Nymphaea stellata Izibo (Zulu) In Zululand (Ubombo Hely- Hutchinson
Willd. = Nymphaea district), tuber boiled (1898), Gammie
nouchali Burm.f. and eaten. In India, (1902), Watt
roots and seeds eaten (1908) and Uphof
Nymphaeaceae Nymphaea tetragona Four Angled Water Lily; Rhizomes used as food Hu (2005)
Georgi Shui Lian (Chinese) in north-west China
Nymphaeaceae Nymphaea tuberosa Tuberous Waterlily, Tubers occasionally Gibbons and
Paine = Nymphaea White Water Lily eaten Tucker (1979),
odorata subsp. tuberosa Fernald et al.
(Paine) Wiersema & (1985), and
Hellq. Facciola (1990)
Nymphaeaceae Ondinea purpurea NF Corms eaten by Les (2003)
Hartog Australian aborigines
Orchidaceae Aceras antropophorum Man Orchid Starchy tubers used for Sezik (2002)
(L.) R.Br. = Orchis making salep flour used
anthropophora (L.) All. in ice-cream
confectionery and
nutritious beverages
Orchidaceae Acianthus apprimus Mountain Gnat Orchid Tubers edible Steenbeeke
D.L. Jones (2001)
Orchidaceae Acianthus collinus NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Acianthus exsertus R.Br Mosquito Orchid Tubers edible Steenbeeke
Orchidaceae Acianthus pusillus Gnat Orchid Tubers edible Steenbeeke
D.L. Jones (2001)
Orchidaceae Anacamptis coriophora NF Starchy tubers used for Ghorbani et al.
(L.) R.M.Bateman making salep flour used (2014) and
in ice-cream Kreziou et al.
production, (2015)
confectionery and
nutritious beverages
Orchidaceae Anacamptis morio (L.) Green-Winged Orchid, As above Kreziou et al.
R.M.Bateman Green-Veined Orchid (2015)
Introduction 9

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Anacamptis morio As above As above Ghorbani et al.
subsp. picta (Loisel.) (2014)
Jacquet & Scappat
Orchidaceae Anacamptis palustris NF As above Ghorbani et al.
(Jacq.) R.M.Bateman, (2014)
Pridgeon & Chase
Orchidaceae Anacamptis Butterfly Orchid As above Kreziou et al.
papilionacea (L.) (2015)
Orchidaceae Anacamptis pyramidalis Pyramidal Orchid As above Sezik (1967),
(L.) Rich. Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010),
Sezik (2002),
Ghorbani et al.
(2014), and
Kreziou et al.
Orchidaceae Arthrochilus huntianus Elbow Orchid Tubers probably edible Steenbeeke
(F.Muell.) Blaxell (2001)
Orchidaceae Barlia robertiana Giant Orchid, Orkida Starchy tubers used for Sezik (2002)
(Loisel.) Greuter Kbira making salep flour used
in ice-cream
confectionery and
nutritious beverages
Ochidaceae Brachycorythis Likos, Ligosi As above Mapunda (2007)
pleistophylla Rchb.f. and Hamisy
Orchidaceae Calendenia caerula Blue Fairy Orchid, Blue Tubers edible Steenbeeke
R.Br. Caladenia (2001)
Orchidaceae Calendenia carnea Pink Fairies, Pink Fringe Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Calendenia filamentosa Daddy Longlegs, Spider Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Calendenia fuscata Dusky Fingers Tubers edible Steenbeeke
(Rchb.f.) M.A.Clem. & (2001)
Orchidaceae Calendenia gracilis Musky Caladenia, Tubers edible Steenbeeke
R.Br. Musky Finger Orchid (2001)
Orchidaceae Calendenia quadrifaria NF Tubers edible Steenbeeke
(R.S.Rogers) D.L.Jones (2001)
Orchidaceae Calendenia tentaculata Green Combed Spider Tubers edible Steenbeeke
Schltdl. Orchid, Fringed Spider (2001)
Orchidaceae Chiloglottis diphylla Common Ant Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Chiloglottis formicifera Ant Orchid Tubers edible Steenbeeke
Fitzg. (2001)
10 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Chiloglottis palachila Clubbed Ant Orchid Tubers edible Steenbeeke
D.L. Jones & (2001)
M.A. Clem.
Orchidaceae Chiloglottis platyptera NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis pluricallata Brown Bird Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis sphyrnoides Ornate Ant Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis Broad-Lip Bird Orchid Tubers edible Steenbeeke
trapeziformis Fitzg. (2001)
Orchidaceae Chiloglottis trilabra NF Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Coelogyne ovalis Lindl. Oval Coelogyne Sotirov (2015)
Orchidaceae Comperia comperiana Komper’s Orchid Starchy tubers used for Sezik (2002)
(Steven) Asch. & making salep flour used
Graebn. = in ice-cream
Himantoglossum production,
comperianum (Steven) confectionery and
P.Delforge nutritious beverages
Orchidaceae Corybas fimbriatus Fringed Helmut Orchid Tubers edible Steenbeeke
(R.Br.) Rchb.f. (2001)
Orchidaceae Corybas hispidus Bristly Helmut Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Corybas montanus Mt Maroon Helmut Tubers edible Steenbeeke
D.L.Jones Orchid (2001)
Orchidaceae Corybas species A Sphagnum Helmut Tubers edible Steenbeeke
Orchid (2001)
Orchidaceae Cryptostylis erecta Tartan Tongue Orchid, Fleshy starchy root Steenbeeke
F. Muell. ex Benth. Turban Orchid, Bonnet eaten (2001)
Orchidaceae Cryptostylis humteriana Leafless Tongue Orchid Fleshy starchy root Steenbeeke
Nicholls eaten (2001)
Orchidaceae Cryptostylis leptochila Small Tongue Orchid, Fleshy starchy root Steenbeeke
R.Br. Red Tongue Orchid eaten (2001)
Orchidaceae Cryptostylis subulata Large Tongue Orchid, Fleshy starchy root Steenbeeke
(Labill.) Rchb.f. Cowslip Orchid eaten (2001)
Orchidaceae Cymbidium Tiger Orchid, Channelled Starch rich pseudobulbs Steenbeeke
canaliculatum R.Br. Cymbidium, Inland Tree eaten cooked or raw (2001)
Orchidaceae Dactylorhiza incarnata Early Marsh Orchid Starchy tubers used for Ghorbani et al.
(L.) Soó making salep flour used (2014)
in ice-cream
confectionery and
nutritious beverages
Orchidaceae Dactylorhiza osmanica The Ottoman Starchy tubers used for Tekinşen and
var. Dactylorhiza making salep flour used Güner (2010) and
osmanica = Dactylorhiza in ice-cream Citil and Tekinşen
osmanica (Klinge) P.F production, (2011)
Hunt & Summerh. confectionery and
nutritious beverages
Introduction 11

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Dactylorhiza romana Roman Dactylorhiza As above Sezik (1967),
(Sebast.) Soó Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010),
and Sezik (2002)
Orchidaceae Dactylorhiza romana Georgian Orchid As above Ghorbani et al.
subsp. georgica (Klinge) (2014)
Soó ex Renz &
Orchidaceae Dactylorhiza saccifera Sack-Carrying As above Kreziou et al.
(Brongn.) Soó Dactylorhiza (2015)
Orchidaceae Dactylorhiza sambucina Elder-Flowered Orchid As above Kreziou et al.
(L.) Soó (2015)
Orchidaceae Dactylorhiza umbrosa Persian Marsh-Orchid As above Ghorbani et al.
(Kar. & Kir.) Wendelbo (2014)
Orchidaceae Dendrobium tarberi Rock Lily, King Orchid
Pseudobulbs eaten Steenbeeke
M.A.Clem. & D.L.Jones baked or raw (2001)
Orchidaceae Dipodium Purple Hyacinth Orchid Fleshy starchy thick Steenbeeke
atropurpureum roots are probably (2001)
D.L.Jones edible
Orchidaceae Dipodium Yellow Hyacinth Orchid Fleshy starchy thick Steenbeeke
hamiltonianum roots are probably (2001)
F.M.Bailey edible
Orchidaceae Dipodium pulchellum Dark Hyacinth Orchid Fleshy starchy thick Steenbeeke
D.L.Jones & M.A.Clem. roots are probably (2001)
Orchidaceae Dipodium punctatum Blotched Hyacinth Fleshy starchy thick Steenbeeke
(Sm.) R.Br. Orchid roots are probably (2001)
Orchidaceae Dipodium roseum Pink Hyacinth Orchid Fleshy starchy thick Steenbeeke
D.L.Jones & roots are probably (2001)
M.A. Clem. edible
Orchidaceae Dipodium variegatum Spotted Hyacinth Orchid Fleshy starchy thick Steenbeeke
M.A.Clem. & D.L.Jones roots are probably (2001)
Orchidaceae Disa engleriana NF Tubers process into a Kasulo et al.
Kraenzl. meatless sausage called (2009)
chikande and Kikande
which is consumed as
relish or as snack
Orchidaceae Disa aequiloba Chikanda Mbozi Tubers edible Nyomoro (2009)
Orchidaceae Disa erubescens Rendle Liseki, Liseku, Makaha Starchy tubers used for Mapunda (2007),
Ga Mlutu, Masekele, making salep flour, Challe and Price
Masekeni, Masekendi, chikande meatless (2009), Nyomoro
Mbozi sausage (2009), and
Hamisy (2010)
Orchidaceae Disa fragrans Schltr. NF Tubers processed into Hamisy (2010)
meatless sausage
12 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Disa hamatopetala Baur’s Disa, Ntetemera Tubers processed into Hamisy (2010)
Rendle = Disa baurii meatless sausage
Bolus kikande
Orchidaceae Disa ochrostachya NF As above Mapunda (2007)
Reichb.f. and Hamisy
Orchidaceae Disa robusta N.E.Br. Chukande Kijike, As above Mapunda (2007),
Likose, Liisek, Manseke, challe and price
Makaha Ga Mlutu, (2009), Kasulo
Masekele, Masekeni et al. (2009), and
Makaha Ga Mlutu, Hamisy (2010)
Masekele, Masekeni
Makaha Ga Mlutu,
Masekele, Masekeni
Orchidaceae Disa tanganyikensis Chikanda Makete Tubers edible Nyomora (2009)
Orchidaceae Disa ukingensis Schltr. NF Tubers processed into Hamisy (2010)
meatless sausage
Orchidaceae Disa walleri Rchb.f. Chikanda Mbeya, Tubers edible Nyomora (2009)
Masekelesekele Njombe
Orchidaceae Disa zombica N.E.Br. NF Tubers processed into Kasulo et al.
meatless sausage (2009) and
kikande Hamisy (2010)
Orchidaceae Diuris abbreviata Lemon Doubletail Fleshy starchy tubers Steenbeeke
F. Muell. ex Benth. edible (2001)
Orchidaceae Diuris alba R.Br. White Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris chrysantha Yellow Donkey Orchid Fleshy starchy tubers Steenbeeke
D.L.Jones & M.A.Clem. edible (2001)
Orchidaceae Diuris dendrobioides Purple Donkey Orchid Fleshy starchy tubers Steenbeeke
Fitzg. edible (2001)
Orchidaceae Diuris goonooensis Western Donkey orchid Fleshy starchy tubers Steenbeeke
Rupp edible (2001)
Orchidaceae Diuris lanceolata Lindl. Golden Moths, Snake Fleshy starchy tubers Steenbeeke
Orchid edible (2001)
Orchidaceae Diuris pedunculata Golden Moths, Small Fleshy starchy tubers Steenbeeke
R.Br. Snake Orchid edible (2001)
Orchidaceae Diuris punctata Sm. Purple Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris semilunata Donkey Orchid, Late Starchy tubers edible Cribb and Cribb
Messmer Leopard Orchid, Spotted (1987), Low
Doubletail Orchid (1989), and Low
Orchidaceae Diuris striata Rupp NF Fleshy starchy tubers Steenbeeke
edible (2001)
Introduction 13

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Diuris sulphurea R.Br Hornet Orchid, Tiger Starchy tubers edible Anonymous
Orchid, Yellow Tiger (2010a, b),
Orchid Anonymous
(2011), Cribb and
Cribb (1987),
Low (1989), Low
(1991), and
Orchidaceae Diuris tricolor Fitzg. Tricolor Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris venosa Rupp Veined Doubletail, Fleshy starchy tubers Steenbeeke
Veined Donkey Orchid, edible (2001)
Goat Orchid
Orchidaceae Epipactis royleana Chhasakrungai Boiled roots eaten Sotirov (2015)
Orchidaceae Eriochilus cucullatus Smooth-Leaf Parson’s Tubers edible Steenbeeke
(Labill.) Rchb.f. Bands, Large (2001)
Orchidaceae Eulophia schweinfurthii Ndulamo, Lisesa Starchy tubers used for Mapunda (2007),
Kraenzl. making salep flour, Challe and Price
chikanda meatless (2009) and
sausage Hamisy (2010)
Orchidaceae Gastrodia procera Large Potato Orchid, Tubers edible Steenbeeke
G.W. Carr Large Cinnamon Bells (2001)
Orchidaceae Gastrodia sesamoides Potato Orchid, Tubers edible Steenbeeke
R.Br. Cinnamon Bells (2001)
Orchidaceae Genoplesium archeri Vriable Midge Orchid Tubers edible Steenbeeke
Orchidaceae Genoplesium filiforme NF Tubers edible Steenbeeke
(Fitzg.) D.L.Jones & (2001)
Orchidaceae Genoplesium fimbriata Fringed Midge Orchid Tubers edible Steenbeeke
(R.Br.) D.L.Jones & (2001)
Orchidaceae Genoplesium Dense Midge Orchid Tubers edible Steenbeeke
nudiscapum (Hook.f.) (2001)
D.L.Jones & M.A.Clem.
Orchidaceae Genoplesium nudum Tiny Midge Orchid Tubers edible Steenbeeke
(Hook.f.) D.L.Jones & (2001)
Orchidaceae Genoplesium pedersonii NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Genoplesium rufum Red Midge Orchid Tubers edible Steenbeeke
(R.Br.) D.L.Jones & (2001)
Orchidaceae Glossodia major R.Br. Large Waxlip Orchid Tubers edible Steenbeeke
Orchidaceae Habenaria adolphii Chinkanda, Tubers edible Nyomora (2009)
Schultr. Vinying’inya, Songea
14 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Habenaria clavata Copper Plant Tubers process into a Kasulo et al.
(Lindl.) Rchb. f. meatless sausage called (2009)
chikande and Kikande
which is consumed as
relish or as snack
Orchidaceae Harbenaria cornuta Chikanda Tubers edible Nyomora (2009)
Orchidaceae Habenaria humilior Chikanda Tubers edible Nyomora (2009)
Orchidaceae Habenaria intermedia Intermediate Habenaria Boiled roots eaten Sotirov (2015)
D. Don
Orchidaceae Habenaria keayi NF Tubers and roots are Menzepoh (2011)
Summerh. used to prepare a food
called “napssié” or
ground meat by the
Bagam tribe in the
subdivision of Galim,
Western region of
Orchidaceae Habenaria praestans Chikanda Mbeya Tubers edible Nyomora (2009)
Orchidaceae Habenaria xanthochlora Mamkumungu, Starchy tubers used for Challe and Price
Schltr. Manseke, making chikanda (2009) and
Mansekemakubwa, meatless sausage Hamisy (2010)
Mviringo, Likose, Liseke
Orchidaceae Habenaria zambesina NF As above Menzepoh (2011)
Orchidaceae Himantoglossum affine NF Starchy tubers used for Sezik (1967),
(Boiss.) Schltr. making salep flour used Sezik and Özer
in ice-cream (1983) cited by
production, Tekinşen and
confectionery and Güner (2010) and
nutritious beverages Ghorbani et al.
Orchidaceae Himantoglossum Komper’s Orchid As above Ghorbani et al.
comperianum (Steven) (2014)
Orchidaceae Malaxix cylindrostachy Cylindric Raceme Boiled roots eaten Sotirov (2015)
(Lindl.) Kuntze Malaxis
Orchidaceae Neotinea maculata Dense-Flowered Orchid Starchy tubers used for Sezik (2002)
(Desf.) Stearn making salep flour used
in ice-cream
confectionery and
nutritious beverages
Orchidaceae Ophrys climacis NF Starchy tubers used for Deniz (2013)
Heimeier & Perschke making salep flour used
in ice-cream
confectionery and
nutritious beverages
Introduction 15

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Ophrys fusca Link Sombre Bee-Orchid, As above Sezik (1967) and
Dark Bee-Orchid Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
Orchidaceae Ophrys holosericea Late Spider Orchid As above Sezik (1967) and
(Burm. f.) Greuter Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
Orchidaceae Ophrys isaura Renz & NF As above Deniz (2013)
Orchidaceae Ophrys lycia Renz & Bee Orchid As above Deniz (2013)
Orchidaceae Ophrys mammosa Desf. Early Spider Orchid As above Tekinşen and
Güner (2010)
Orchidaceae Ophrys phaseliana Sawfly Orchid As above Deniz (2013)
D.Rückbr. & U.Rückbr
Orchidaceae Ophrys scolopax Cav. Woodcock Bee-Orchid, As above Ghorbani et al.
Woodcock Orchid (2014)
Orchidaceae Ophrys sphegodes Mill. Early Spider Orchid As above Ghorbani et al.
Orchidaceae Ophrys sphegodes NF As above Ghorbani et al.
subsp. mammosa (Desf.) (2014)
Soó ex E.Nelson
Orchidaceae Orchis adenocheila NF As above Ghorbani et al.
Czerniak (2014)
Orchidaceae Orchis anatolica Boiss. Anatolian Orchid As above Tekinşen and
Güner (2010) and
Citil and Tekinşen
Orchidaceae Orchis anthropophora Man Orchid As above Kreziou et al.
(L.) All. (2015)
Orchidaceae Orchis conopea Fragrant Orchid As above Grieve (1971)
Gras = Gymnadenia
conopsea (L.) R.Br.
Orchidaceae Orchis coriophora L. = Fragrant Orchid As above Grieve (1971),
Anacamptis coriophora Tekinşen and
(L.) R.M. Bateman, Güner (2010),
Pridgeon & MW Chase and Citil and
Tekinşen (2011)
Orchidaceae Orchis italica Poir Naked Man Orchid, As above Tekinşen and
Italian Orchid Güner (2010),
Citil and Tekinşen
(2011), and
Kreziou et al.
16 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Orchis latifolia L. = Early Marsh Orchid As above Grieve (1971)
Dactylorhiza incarnata
(L.) Soó
Orchidaceae Orchis longicruris Naked Man Orchid As above Grieve (1971)
Link = Orchis italica
Orchidaceae Orchis maculata L. = Heath Spotted Orchid As above Grieve (1971)
Dactylorhiza maculata
(L.) Soó
Orchidaceae Orchis mascula (L.)L. Early Purple Orchid Starchy tubers used for Ghorbani et al.
salep production (2014)
Orchidaceae Orchis mascula subsp. Early Purple Orchid As above Sezik (1967),
pinetorum (Boiss. & Sezik and Özer
Kotschy) (1983) cited by
E.G.Camus = Orchis Tekinşen and
mascula subsp. mascula Güner (2010),
Grieve (1971),
Hawkes (1944),
and Kreziou et al.
Orchidaceae Orchis militaris L. Military Orchid As above Hawkes (1944),
Grieve (1971),
and Kreziou et al.
Orchidaceae Orchis morio Green-Winged Orchid As above Hawkes 1944,
L = Anacamptis morio Green-Veined Orchid Grieve 1971, and
(L.) R.M. Bateman, Tekinşen and
Pridgeon & MW Chase Güner (2010),
Citil and Tekinşen
Orchidaceae Orchis palustris Jacq. = Toothed Orchid, As above Tekinşen and
Anacamptis palustris Three-Toothed Orchid Güner (2010) and
(Jacq.) R.M. Bateman, Citil and Tekinşen
Pridgeon &MW Chase (2011)
Orchidaceae Orchis provincialis Provence Orchid, Orchis As above Kreziou et al.
Balbis ex Lamarck & De Provence (2015)
Orchidaceae Orchis pyramidalis L. = Pyramidal Orchid As above Grieve (1971)
Anacamptis pyramidalis
Orchidaceae Orchis saccifera Brong. NF As above Grieve (1971)
= Dactylorhiza
saccifera (Brongn.) Soó.
Orchidaceae Orchis simia Lam Monkey Orchid As above Tekinşen and
Güner (2010) and
Ghorbani et al.
Orchidaceae Orchis spitzelii Saut. ex L’orchis De Spitzel As above Sezik (1967),
W.D.J.Koch Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
Introduction 17

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Orchis tridentata Scop. Three-Toothed Orchid Starchy tubers used for Tekinşen and
= Neotinea tridentata making salep flour used Güner (2010) and
(Scop.) R.M. Bateman, in ice-cream Citil and Tekinşen
Pridgeon MW Chase production, (2011)
confectionery and
nutritious beverages
Orchidaceae Orchis ustulata L. = Burnt-Tip Orchid As above Grieve (1971)
Neotinea ustulata (L.)
R.M.Bateman, Pridgeon
& M.W.Chase.
Orchidaceae Orthoceras strictum Horned Orchid, Birds’s Tuberoids probably Steenbeeke
R.Br. Mouth Orchid edible (2001)
Orchidaceae Peristylus constrictus Constricted Peristylus Tubers eaten Sotirov (2015)
Lindley) Lindley
Orchidaceae Platanthera calvigera Club Carrying Boiled roots eaten Sotirov (2015)
Lindl Platanthera
Orchidaceae Platanthera bifolia (L.)Lesser Butterefly Orchid Starchy tubers used for Ghorbani et al.
L.C.Rich making salep (2014)
Orchidaceae Prasophyllum brevilabre Short-Lipped Leek Tubers edible Steenbeeke
(Lindl.) Hook.f. Orchid (2001)
Orchidaceae Prasophyllum campestre Starry Leek Orchid Tubers edible Steenbeeke
R.J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum Ben Lomond Leek Tubers edible Steenbeeke
caudiculum D.L. Jones Orchid (2001)
Orchidaceae Prasophyllum dossenum NF Tubers edible Steenbeeke
J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum elatum Tall Leek Orchid, Piano Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Prasophyllum flavum Yellow leek Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Prasophyllum odoratum Scented Leek Orchid, Tubers edible Steenbeeke
R.S.Rogers Sweet Leek Orchid (2001)
Orchidaceae Prasophyllum patens Broad-Lipped Leek Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Prasophyllum rogersii Marsh Leek Orchid Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Prasophyllum solstitium Summer Leek Orchid Tubers edible Steenbeeke
R.J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum species A Tablelands Leek Orchid Tubers edible Steenbeeke
Orchidaceae Pteroglossaspis Mbozi As above Nyomora (2009)
eustachya Rchb.f.
Orchidaceae Pterostylis abrupta Abrupt Greenhood, Tubers edible Steenbeeke
D.L.Jones Drooping Greenhood (2001)
Orchidaceae Pterostylis alata NF Tubers edible Steenbeeke 2001
(Labill.) Rchb.f.
Orchidaceae Pterostylis bicolor Two-Colour Greenhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
18 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Pterostylis boormanii Boorman’s Greenhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
Orchidaceae Pterostylis chaetophora Hair-Lip Ruddyhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
Orchidaceae Pterostylis coccina Alpen Greenhood Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Pterostylis curta R. Br Blunt Greenhood Tubers edible Anonymous
(2010a, b) and
Orchidaceae Pterostylis cycnocephala Swan Greenhood Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Pterostylis daintreana NF Tubers edible Steenbeeke
F.Muell. ex Benth. (2001)
Orchidaceae Pterostylis decurva Summer Greenhood Tubers edible Steenbeeke
R.S. Rogers (2001)
Orchidaceae Pterostylis fischii NF Tubers edible Steenbeeke
Nicholls (2001)
Orchidaceae Pterostylis hamata NF Tubers edible Steenbeeke
Blackmore & Clemesha (2001)
Orchidaceae Pterostylis hildae Rainforest Greenhood Tubers edible Steenbeeke
Nicholls (2001)
Orchidaceae Pterostylis laxa NF Tubers edible Steenbeeke
Blackmore (2001)
Orchidaceae Pterostylis longicurva NF Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Pterostylis longifolia Tall Greenhood Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Pterostylis longipetala NF Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Pterostylis mutica R.Br. Midget Greenhood Tubers edible Steenbeeke
Orchidaceae Pterostylis nutans R.Br. Nodding Greenhood Tubers edible Steenbeeke
Orchidaceae Pterostylis obtusa R.Br. Jug-Lip Greenhood Tubers edible Steenbeeke
Orchidaceae Pterostylis parviflora Tiny Greenhood, Jug Tubers edible Steenbeeke
R.Br. Orchid, Green Snail (2001)
Orchidaceae Pterostylis pedunculata Maroonhood Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Pterostylis praetermissa NF Tubers edible Steenbeeke
M.A.Clem. & D.L.Jones (2001)
Orchidaceae Pterostylis reflexa R.Br. NF Tubers edible Steenbeeke
Orchidaceae Pterostylis setifera NF Tubers edible Steenbeeke
M.A. Clem. (2001)
Orchidaceae Pterostylis species B NF Tubers edible Steenbeeke
Introduction 19

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Pterostylis species C NF Tubers edible Steenbeeke
Orchidaceae Pterostylis species D NF Tubers edible Steenbeeke
Orchidaceae Pterostylis truncate Little Dumpies, Sasauge Tubers edible Steenbeeke
Fitzg. Greenhood (2001)
Orchidaceae Pterostylis woollsii Long-Tailed Greenhood, Tubers edible Steenbeeke
Fitzg. Chinaman Greenhood (2001)
Orchidaceae Roeperocharis Kaloba, Masekel Tubers process into a Challe and Price
wentzeliana Kraenzl meatless sausage called (2009) and
chikande Hamisy (2010)
Orchidaceae Satyrium acutirostrum Chikande As above Hamisy (2010)
Orchidaceae Satyrium amblyosaccos NF Tubers process into a Kasulo et al.
Schltr. meatless sausage called (2009)
chikande and kikande,
which is consumed as
relish or as snack
Orchidaceae Satyrium atherstonei Chikande Ligos, Ingingi, Starchy tubers used for Mapunda (2007),
Rchb.f. = Satyrium Jike, Lidala, Lisekejike, making salep flour, Challe and Price
trinerve Lindl. Lisekeni, Lisekenilidala, chikanda meatless (2009) and
Madala, sausage Hamisy (2010)
Numbunumbu, Sidala,
Visekenividala, Vijike
Orchidaceae Satyrium breve Rolfe Chikande Tubers edible Nyomora (2009)
Orchidaceae Satyrium buchananii Dochamua, Ligosi, Tubers process into a Mapunda (2007),
Schltr. Likosi, Lisekedochamua, meatless sausage called Kasulo et al.
Lisekedume, chikande and (2009) and
Lisekekiume, Magosi, kikande,which is Hamisy (2010)
Masekenidume, consumed as relish or
Masekeni magosi, as snack, starchy tubers
Sisekeni sigosi, used for making salep
Titisigosi, flour
Orchidaceae Satyrium carsonii Rolfe NF Tubers process into a Kasulo et al.
meatless sausage called (2009)
chikande and kikande,
which is consumed as
relish or as snack
Orchidaceae Satyrium chlorocorys Chikanda Jike Rukwa Tubers edible Nyomora (2009)
Orchidaceae Satyrium crassicaule Chikanda, Kikande, Tubers process into a Nyomora (2009)
Rendle Makete, Simbegi meatless sausage called and Hamisy
chikande and kikande, (2010)
which is consumed as
relish or as snack
Orchidaceae Satyrium nepalense Nepal Satyrium Pseudobulbs eaten Sotirov (2015)
D.Don boiled
Orchidaceae Satyrium robustum Kidume Tubers processed into Hamisy (2010)
Schltr. meatless sausage
20 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Satyrium sacculatum Chikanda, Kikande, Tubers edible Nyomora (2009)
(Rendle) Rolfe Masehelesehele, Vikan,
Dokando, Songea
Orchidaceae Satyrium volkensii Volkens ‘Satyrium Tubers processed into Hamisy (2010)
Schltr. meatless sausage
Orchidaceae Serapias vomeracea ssp. NF Starchy tubers used for Tekinşen and
orientalis making salep Güner (2010),
Grueter = Serapias Citil and Tekinşen
orientalis (Greuter) (2011), Sezik
H. Baumann & Künkele (1967), Sezik and
Özer (1983) cited
by Tekinşen and
Güner (2010)
Orchidaceae Steveniella satyrioides NF Starchy tubers used for Ghorbani et al.
(Spreng.) Schltr making salep (2014)
Orchidaceae Thelymitra carnea R.Br. Tiny Sun Orchid, Pink Tubers edible Steenbeeke
Sun Orchid (2001)
Orchidaceae Thelymitra circumsepta Naked Sun Orchid Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Thelymitra cyanea Veined Sun Orchid Tubers edible Steenbeeke
(Lindl.) Benth. (2001)
Orchidaceae Thelymitra fragrans NF Tubers edible Steenbeeke
D.L.Jones & M.A.Clem. (2001)
Orchidaceae Thelymitra Scented Sun Orchid Tubers edible Steenbeeke
megacalyptra Fitzg. (2001)
Orchidaceae Thelymitra nuda R.Br. Plain Sun Orchid Tubers edible Steenbeeke
Orchidaceae Thelymitra pauciflora Slender sun Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Tipularia discolour Crane Fly Orchid, Starchy corm eaten Sotirov (2015)
(Pursh) Nuttall Crippled Cranefly Orchid
Oxalidaceae Oxalis corymbosa DC. = Pink Shamrock Sweet crisp swollen tap Cribb and Cribb
Oxalis debilis var. root eaten (1987)
corymbosa (DC.)
Oxalidaceae Oxalis deppei Lodd. ex Orach, Deppei Wood Fleshy root eaten Fernald et al.
Sweet Sorrel boiled in Mexico (1985) and
Facciola (1990)
Oxalidaceae Oxalis perennans Haw. Grassland Wood Sorrel Small tap roots eaten Low (1991)
by aborigines
Oxalidaceae Oxalis radicosa A.Rich. Dwarf Wood Sorrel Small tap roots eaten Low (1991)
= Oxalis corniculata L. by aborigines
Oxalidaceae Oxalis sp. Wood Sorrel Small tap roots eaten Low (1991)
by aborigines
Oxalidaceae Oxalis stricta L. Yellow Wood Sorrel Roots edible Yanovsky (1936),
Uphof (1968),
and Facciola
Introduction 21

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Oxalidaceae Oxalis tuberosa Molina Oca Tubers eaten boiled, Facciola (1990),
roasted or candied, one Groen et al.
of the principal food (1996), Flores
crop of the Indians in et al. (2003), van
the Andes, second to Wyk (2006), and
potato, tubers can be Santich et al.
eaten unpeeled, raw, in (2008),
salads, pickled, boiled,
fried or in soups and
stews. Also can be
dried in sun to sweeten
Oxalidaceae Oxalis violacea L. = Violet Wood Sorrel Roots edible Yanovsky (1936)
Oxalis debilis var. and Facciola
corymbosa (DC.) (1990)
Piperaceae Piper methysticum L.f. Kava, Kava Kava, Awa, The root and rhizome McDonald and
Kava Pepper (underground stem) of Jowitt (2000) and
kava are used to NCAM (2006)
prepare beverages,
extracts, capsules,
tablets and topical
solutions. Kava is
widely and commonly
consumed as a social
beverage to establish
kinship in the Pacific
island communities
Poaceae Agropyron repens (L.) English Couch, Quick Roots ground into a Cribb and Cribb
P. Beauv. = Elymus Grass meal and used to make (1987)
repens (L.) Gould bread
Poaceae Bambusa pallida Munro Mokal Bah (Assamese); Young rhizomes are Patiri and Borah
Mai Phiu, Phai Song eaten as vegetable after (2007)
Kham (Thai) processing
Poaceae Bambusa tulda Roxb. Spineless Indian As above Patiri and Borah
Bamboo, Calcutta Cane; (2007)
Jati Bah (Assamese)
Poaceae Chrysopogon Vetiver Grass, Khas- Vetiver is used Balasankar et al.
zizanioides (L.) Roberty Kahs, Khus Grass domestically in (2013)
cooking; it’s infused in
tea and also used in
Poaceae Eleusine indica (L.) Indian Goosegrass, Roots edible raw Kunkel (1984)
Gaertn. Wiregrass,
Poaceae Hordeum bulbosum L. Abu Suwaif Bulbous roots chewed Tanaka (1976)
and eaten occasionally and Facciola
22 Introduction

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Poaceae Imperata cylindrica L. Blady Grass, Alang- Aborigine children Burkill (1966),
Alang, Lalang Wooly sucked the roots, Cribb and Cribb
Grass underground shoots (1987), Low
like sugarcane, used as (1991), and Hu
survival food, a kind of (2005)
beer is made from the
roots in Peninsular
Malaysia China: fresh
rhizome chewed by
rural people for the
sweet juice
Poaceae Phragmites australis Common Reed Grass Rhizomes coked like Tanaka (1976),
(Cav.) Trin. Ex Steud. potatoes Fernald et al.
(1985), and
Facciola (1990)
Poaceae Saccharum spontaneum Wild Cane, Wild Young shoots and Patiri and Borah
L. Sugarcane, Fodder Cane; rhizomes are eaten as (2007)
Khagori (Assamese); sugarcane, sweet in
Mojora (Mishing) taste
Poaceae Triticum repens L. = Couch Grass, Twitch, In Norway, roots Sayce (1953)
Elymus repens (L.) Quick Grass, Quitch ground into flour
Gould. Grass, Dog Grass
Poaceae Vetiveria nigritana Black Vetiver grass; Nigeria (Kano State, Mortimore (1989)
(Benth.) Jema (Huasa) northern): roots eaten.
Stapf = Chrysopogon
nigritanus (Benth.)
Poaceae Vetiveria zizanioides Vetiver Grass, Khas- Roots used to prepare Pareek and
(L.) Nash = Chrysopogon Khas Grass, Akar Wangi sherbets or soft drinks Kumar (2013)
zizanioides (L.) Roberty. (Malay) during summer in and Balasankar
Nothern India. Vetiver et al. (2013)
oil is used for
flavouring syrups,
ice-cream and food
preservation. Khus
essence is used in cool
drinks, for reducing
pungency of chewing
tobacco preparations
and to provide sweet
note to other
Vetiver roots is used
domestically in
cooking; it’s infused in
tea and also used in
Poaceae Zizaniopsis miliacea Water Millet, Southern Young rhizome, cut Gibbons and
(Michx.) Döll & Asch. Wild Rice into pieces cooked and Tucker (1979),
served with butter Fernald et al.
(1985), and
Facciola (1990)
Introduction 23

Table 1 (continued)
Family Scientific Name Names Edible Part Use Reference
Rubiaceae Ixora subsessilis Wall. Dieng Jowat Flower, shoot, root Sawian et al.
ex G.Don (Meghalaya); Nang Guo eaten in Meghalaya (2007)
Long Chuan Hua
Rubiaceae Morinda officinalis F.C. Chinese Herbal Morinda, In China, fresh or dried Hu (2005), Li et
How Medicinal Indian roots are cooked with al. (2009)
Mulberry, Morinda Root pork for a broth as
health food called
Chinese bupin (Hu
2005). In South China,
Hong Kong and Macao,
this plant has been
developed into various
health foods, such as
‘Ba-ji-tian wine’,
‘Ba-ji-zi-bu Gao’ (Li
et al. 2009)
Rubiaceae Paederia foetida L. Rekang Nemthu Roots eaten in Karbi Kar and
Assam Borthakur (2008)
Rubiaceae Paederia stenobotrya White Paedaria Root ground with Hu (2005)
Merr. soaked soybean for a
milk, cooked and given
to people with jaundice
Simaroubaceae Eurycoma longifolia Tongkat Ali (Malay) Traditional users of Low et al. (2013)
Jack Tongkat Ali brew tea
from the dried chips of
the tree’s root. Tea,
coffee and carbonated
beverages, pre-mixed
with the root extract are
available commercially
for the improvement of
general health and
libido in Malaysia
NF not found

is processed into nutritious and wholesome from the genera Disa, Habenaria and Satyrium.
beverages, confectionary and ice-cream. Salep is Many species in the orchid genera Pterostylis,
especially popular in Middle East and some parts Diuris and Thelymitra have edible tubers. Other
of Europe. The major salep producing orchid orchid species with edible roots and tubers are
species are from the genera Anacamptis, Orchis, listed in Table 1.
Ophrys, Disa, Dactylorhiza and Himantoglossum.
In Africa, edible orchid tubers are collected from
the wild and processed into a meatless sausage References
locally called chinaka, chikande and kikande,
which is consumed as relish or just as a snack Altschul SVR (1973) Drugs and foods from little-known
plants. Harvard University Press, Cambridge, MA,
(Kasulo et al. 2009). The tubers are also used in
518 pp
soups which are reported to be served in some Anonymous (2010a) Diuris sulphurea. http://www.yarra-
international tourist hotels (Hamisy 2010). The
main orchid species used for this purpose are Directory/Lower_Storey/Orchids/Diuris_sulphurea
24 Introduction

Anonymous (2010b) Pterostylis curta. http://www.yarra- Flores HE, Walker TS, Guimaraes RL, Bais HP, Vivanco JM (2003) Andean root and tuber crops: underground
Directory/Lower_Storey/Orchids/Pterostylis_curta rainbows. HortSci 38(2):161–167
Anonymous (2011) Diuris sulphurea (tiger orchid). http:// Fox FW, Young MMN, Hallowes D (1982) Food from the veld: edible wild plants of southern Africa botanically
php?1502-Diuris-sulphurea-%28Tiger-Orchid%29 identified and described. Delta Books, Cape Town
Balasankar D, Vanilarasu K, Preetha PS, Umadevi SRM, 399pp
Bhowmik D (2013) Traditional and medicinal uses of Freedman RL (2009) Famine foods database. Purdue
vetiver. J Med Plant Stud 1(3):191–200 University.
Baranov AI (1967) Wild vegetables in Manchuria. Econ FamineFoods/ff_home.html
Bot 21:140–155 Gammie GA (1902) A note on plants used for food during
Bender DA (2009) A dictionary of food and nutrition, 3rd famines and seasons of scarcity in the Bombay
edn. Oxford University Press, Oxford Presidency. India Bot Surv Rec 2(2):171–196
Bermejo JEH, León J (eds) (1994) Neglected crops: 1492 Ghorbani A, Gravendeel B, Naghibi F, de Boer H (2014)
from a different perspective, Plant production and pro- Wild orchid tuber collection in Iran: a wake-up call for
tection series no. 26. FAO, Rome, pp 205–209 conservation. Biodivers Conserv 23(11):2749–2760
Burkill IH (1966) A dictionary of the economic products Gibbons E, Tucker G (1979) Euell Gibbons’ handbook of
of the Malay Peninsula. Revised reprint, 2 volumes. edible wild plants. The Doning Co, Virginia Beach
Ministry of Agriculture and Co-operatives, Kuala Grieve M (1971) A modern herbal. Penguin, 2 vols. Dover
Lumpur, Malaysia, vol 1 (A–H) pp 1–1240, vol 2 Publications, New York, 919 pp
(I–Z). pp 1241–2444 Groen LE, Siemonsma JS, Jansen PCM (1996) Minor
Challe JF, Price LL (2009) Endangered edible orchids and species yielding non-seed carbohydrates. In: Flach M,
vulnerable gatherers in the context of HIV/AIDS in Rumawas F (eds) Plants yielding non-seed carbohy-
the Southern Highlands of Tanzania. J Ethnobiol drates, Plant resources of South East Asia no. 9. Prosea
Ethnomed 5:41 Foundation, Bogor, pp 165–186
Chase AW (1900) Dr. Chase’s recipes; or, information for Hamisy WC (2010) Development of conservation strate-
everybody: an invaluable collection of about eight gies for the wild edible orchids in Tanzania. Progress
hundred practical recipes, 40th edn. R. A. Beal report for the Rufford Small Grant Foundation
Publisher, Ann Arbor, 648 pp December 2010, Arusha, Tanzania
Citil OB, Tekinsen KK (2011) A comparative study on Harden GJ (ed) (1990) Flora of New South Wales, vol 1.
fatty acid composition of salep obtained from New South Wales University Press, Sydney, 601 pp
some Orchidaceae species. Chem Nat Comp Hawkes AD (1944) Economic importance of the
46(6):943–945 Orchidaceae. II. Amer Orchid Soc Bull 13:56–58
Codex (2014) Draft revision of the classification of foods Hedrick UP (1972) Sturtevant’s edible plants of the world.
and animal feeds: selected vegetable commodity Dover Publications, New York, 686 pp
groups. CX/PR 14/46/7. Hely-Hutchinson WF (1898) Famine plants in Zululand.
ings/ccpr/ccpr46/pr46_07e.pdf Great Britain. Royal Botanic Gardens at Kew. Bull
Coffey T (1994) The history and folklore of North Misc Inf 135:52–54
American wildflowers. Houghton Mifflin Harcourt, Hu SY (2005) Food plants of China. The Chinese
Boston, 356pp University Press, Hong Kong, 844 pp
Cribb AB, Cribb JW (1987) Wild food in Australia, 2nd Irvine FR (1952) Supplementary and emergency food
edn. Fontana Collins, Sydney, 240 pp plants of West Africa. Econ Bot 6(1):23–40
Dalziel JM (1955) The useful plants of west tropical Africa Jansen PCM (1996) Plectranthus rotundifolius (Poiret)
(Reprint of 1937 ed.) Crown Agents for Overseas Sprengel. In: Flach M, Rumawas F (eds) Plants yield-
Governments and Administrations, London, 612 pp ing non-seed carbohydrates, Plant resources of South
den Bergh V (1996) Stachys sieboldii Miquel. In: Flach East Asia no. 9. Prosea Foundation, Bogor,
M, Rumawas F (eds) Plants yielding non-seed carbo- pp 141–143
hydrates, Plant resources of South East Asia no. 9. Kar A, Borthakur SK (2008) Wild vegetables of Krabi –
Prosea Foundation, Bogor, pp 155–156 Anglong district, Assam. Nat Prod Rad 7(5):448–460
Deniz IG (2013) Critically endangered (cr) orchid species Kasulo V, Mwabumba L, Cry M (2009) A review of edible
of Antalya Province of Turkey. Acta Hortic orchids in Malawi. J Hort For 1(7):133–139
1002:37–41 Kreziou A, de Boer H, Gravendeel B (2015) Harvesting of
Facciola S (1990) Cornucopia. A source book of edible salep orchids in north-western Greece continues to
plants. Kampong Publications, Vista, 677 pp threaten natural populations. Oryx 2015:1–4
Fairchild D (1930) Exploring for plants. Macmillan, Kunkel G (1984) Plants for human consumption. An
New York annotated checklist of the edible phanerogams and
Fernald ML, Kinsey AC, Rollins RC (1985) Edible wild ferns. Koeltz Scientific Books, Koenigstein
plants of eastern North America, 2nd edn. Harper & Les DH (2003) Nymphaeales. Wiley, Chichester. doi:
Row, New York 10.1038/npg.els.0003697.
Introduction 25

Li N, Qin LP, Han T, Wu YB, Zhang QY, Zhang H (2009) Patiri B, Borah A (2007) Wild edible plants of Assam.
Inhibitory effects of Morinda officinalis extract on bone Geetakhi Printers and Publishers, Guwahati
loss in ovariectomized rats. Molecules 14(6):2049–2061 Paton DN, Dunlop JC (1904) The nutritive values of some
Lim TK (2014) Edible medicinal and non medicinal uncultivated foods used by the Bhils during recent
plants volume 8, flowers. Springer, Dordrecht/ famines. Agric Ledger 6:37–73
Heidelberg/New York Phillips R, Rix M (1993) Vegetables. Pan Books, London,
Low T (1989) Bush tucker – Australia’s wild food harvest. 270pp
Angus & Robertson, North Ryde, 233 pp Pongpangan S, Poobrasert S (1985) Edible and poisonous
Low T (1991) Wild food plants of Australia. Angus & plants in Thai forests. Science Society of Thailand,
Robertson, North Ryde, 240 pp Science Teachers Section, Bangkok, 206 pp
Low BS, Choi SB, Abdul Wahab H, Kumar Das P, Chan Popenoe H, King SR, Léon J, Kalinkowski LS, Vietmeyer
KL (2013) Eurycomanone, the major quassinoid in ND, Dafforn M (1989) Lost crops of the Incas. Little
Eurycoma longifolia root extract increases spermato- known plants of the Andes with promise for world-
genesis by inhibiting the activity of phosphodiesterase wide cultivation. National Academy Press,
and aromatase in steroidogenesis. J Ethnopharmacol Washington, pp 37–46
149(1):201–207 Read BE (ed) (1946) Famine foods listed in the Chiu
Mapunda LND (2007) Edible orchids in makete district, huang pen ts’ao [of Ting Wang Chou]: giving their
the southern highlands of Tanzania: distribution, pop- identity, nutritional values and notes on their prepara-
ulation and status. Masters thesis Swedish Biodiversity tion. Henry Lester Institute of Medical Research,
Centre Uppsala, Sweden Shanghai, 93pp
McDonald D, Jowitt A (2000) Kava in the Pacific Islands: Santich B, Bryant G, Bacon J, Clifton C, Connor D,
a contemporary drug of abuse? Drug Alcohol Rev Felder A, Foster S, Graue J, Loyer J, Morrachian
19(2):217–227 M, Raab C, Sandall P, Santich S, Stybe K, Stybe R
Medhi P, Borthakur SK (2012) Phytoresources from north (2008) Edible: The illustrated guide to the world’s
Cachar Hills of Assam – III: edible plants sold at food plants. Cameron House, South Australia,
Haflong market. Indian J Nat Prod Resour 3(1):84–109 360 pp
Menzepoh SB (2011) Les orchidées comestibles chez le Saunders CF (1920) Useful wild plants of the United
peuple Bagam, au Cameroun. Biotechnol Agron Soc States and Canada. Robert M. Mcbride & Co,
Environ 15:4 New York
Mortimore M (1989) Adapting to drought. Farmers, fam- Sawian JT, Jeeva S, Lyndem FG, Mishra BP, Laloo RC
ines and desertification in West Africa. Cambridge (2007) Wild edible plants of Meghalaya, North-east
University Press, Cambridge, 324 pp India. Nat Prod Rad 6(5):410–462
Morton JF (1976) Herbs and spices. Golden Press, Sayce RU (1953) Need years and need foods.
New York, 160pp Montgomeryshire Collections 53(Part 1):55–80
National Centre for Complementary and Alternative Schofield J (2003) Discovering wild plants: Alaska,
Medicine (NCAM) (2006) Kava. US Department of Western Canada, the Northwest, First edn. Alaska
Health and Human Services. Northwest Books, Anchorage, 353 pp
sites/ Sezik E (1967) Türkiye’nin salepgilleri ticari salep
Kava_06-15-2012_0.pdf çeşitleri ve özellikle Muğla salebi üzerinde
Nyomora AMS (2009) Distribution and abundance of the araştırmalar. Doktora Tezi. No: 34. Istanbul
edible orchids of the southern highlands of Tanzania. Sezik E (2002) Destruction and conservation of Turkish
Tanzania J Sci 31(1):45–54 orchids. In: Sener B (ed) Biodiversity: biomolecular
Ochse JJ, van den Brink RCB (1980) Vegetables of the aspects of biodiversity and innovative utilization.
Dutch Indies, 3rd edn. Ascher & Co., Amsterdam, Springer, New York, pp 391–400
1016 pp Sezik E, Özer B (1983) Kastamonu salebinin menşei ve
Ong HC (1996) Nelumbo nucifera Gaertner. In: Flach M, Kastamonu civarının orkideleri. Türkiye Bilimsel ve
Rumawas F (eds) Plant resources of South East Asia Teknik Araştırma Kurumu. Proje No: TBAG-424.
No. 9. Plants yielding non-seed carbohydrates. Prosea Ankara
Foundation, Bogor, pp 131–133 Shortt J (1887–1888) List of wild plants and vegetables
Pareek A, Kumar A (2013) Ethnobotanical and pharma- used as food by people in famine times. Indian Forester
ceutical uses of Vetiveria zizanioides (Linn) Nash: a 3:232–238
medicinal plant of Rajasthan. Int J Life Sci Pharm Sci Sotirov KZ (2015) Edible orchids. http://www.ionopsis.
50:L12–L18 com/edible_orchids.htm
Parmentier A (1781) Recherches sur les végétaux nourri- Steenbeeke G (2001) List of plant species from northern
sans qui, dans les temps de disette, peuvent remplacer NSW that may be used as food plants. http://brg.cma.
les alimens ordinaires. Avec de nouvelles observations
su la culture des pommes de terre. Imprimerie Royale, Surhone LM, Tennoe MT, Henssonow SF (2011) Orris
Paris, 599 pp root. Betascript Publishing, Beau Basin, 100 pp
26 Introduction

Tanaka T (1976) Tanaka’s cyclopaedia of edible plants of Van Wyk BE (2006) Food plants of the world: an illus-
the world. Keigaku Publishing, Tokyo, 924 pp trated guide. Timber Press, 480 pp
Tekinşen KK, Güner A (2010) Chemical composition and Villamayor FG Jr, Jukema J (1996) Maranta arundinacea
physicochemical properties of tubera salep produced L. In: Flach M, Rumawas F (eds) Plants yielding non-
from some Orchidaceae species. Food Chem seed carbohydrates, Plant resources of South East Asia
121:468–471 no. 9. Prosea Foundation, Bogor, pp 113–116
The Plant List (2013) Version 1.1. Published on the Watt G (1908) The commercial products of India:
Internet; being an abridgement of “the dictionary of the eco-
Uphof JCT (1968) Dictionary of economic plants. Verlag nomic products of India”. J. Murray, London,
von Cremer, Lehrer 1189 pp
Van den Bergh MH (1994) Minor vegetables. In: Yanovsky E (1936) Food plants of the North American
Siemonsma JS, Piluek K (eds) Vegetables, Plant Indians. United States Department of Agriculture.
resources of South East Asia no. 8. Prosea Foundation, Miscellaneous Publication. No. 237. U.S. Department
Bogor, pp 280–310 of Agriculture, Washington, DC
Iris x germanica

Scientific Name ana Prodán, Iris × officinalis Salisb., Iris × pal-

lida Ten. (Illeg.), Iris pallida subsp. australis
Iris x germanica L. (Tod.) K.Richt., Iris × piatrae Prodán, Iris × red-
outeana Spach, Iris × repanda Berg, Iris × roth-
schildii Degen, Iris × sambucina L.,
Synonyms Iris × spectabilis Salisb., Iris × squalens L.,
Iris × squalens var. biflora Prodán & Buia,
Iris × alba Savi, Iris × amoena DC., Iris × atrovio- Iris × squalens var. rosea Prodán & Buia,
lacea Lange, Iris × australis Tod., Iris × belouinii Iris × superba Berg, Iris × tardiflora Berg,
Bois & Cornuault, Iris × biliottii Foster, Iris × bui- Iris × trojana A.Kern. ex Stapf, Iris × varbossa-
ana Prodán, Iris × buiana var. virescens Prodán, nia K.Malý, Iris variegata var. lurida (Aiton)
Iris × croatica Prodán, Iris × croatica Horvat & Nyman, Iris × venusta J.Booth ex Berg,
M.D.Horvat (Illeg.), Iris × cypriana Foster & Iris × violacea Savi, Iris × vulgaris Pohl
Baker, Iris × deflexa Knowles & Westc., Iris × flo-
rentina L., Iris × florentina var. pallida Nyman,
Iris × florentinoides Prodán ex Nyar., Iris × ger- Family
manica var. alba Dykes, Iris × germanica var.
amas Dykes, Iris × germanica var. askabadensis Iridaceae
Dykes, Iris × germanica var. australis (Tod.) Dykes
Iris × germanica var. florentina (L.) Dykes,
Iris × germanica var. fontarabie Dykes, Common/English Names
Iris × germanica var. gypsea Rodigas, Iris × ger-
manica var. kharput Dykes, Iris × germanica Bearded Iris, Common Orrisroot, Flag, Florentine
var. lurida (Aiton) Nyman, Iris × germanica var. Orris, Florentine Iris, Garden Iris, German Iris,
nepalensis (Wall. ex Lindl.) Herb., Iris × ger- German Orrisroot, Iris, Orris, Orrisroot, Purple
manica var. sivas G.Nicholson, Iris × humei Flag, Queen Elizabeth Root, Tall Bearded
G.Don, Iris × laciniata Berg, Iris × latifolia German Iris, Tall Bearded Iris
Gilib. (Inval.), Iris × lurida Aiton, Iris × mac-
rantha Simonet, Iris × mesopotamica Dykes,
Iris × murorum Gaterau, Iris × neglecta Vernacular Names
Hornem., Iris × nepalensis Wall. ex Lindl.,
Iris × nostras Garsault (Inval.), Iris × nyarady- Arabic: Irsa

© Springer International Publishing Switzerland 2016 27

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_2
28 Iridaceae

Chinese: Déguó Yuānwěi tile hybrid between I. pallida and I. variegata L.,
Czech: Kosatec Německý both of which also have the chromosome number
Danish: Have-Iris, Iris, Sværdlilie 2n = 24 (Henderson 1992, 2002).
Dutch: Blauwe Lis, Duitse Lis, Duitse Lis Sort,
Lis Sort
Esperanto: Irido Ĝardena, Irido Germana Agroecology
Estonian: Aediiris
Finnish: Saksankurjenmiekka, Sininen Iris x germanica thrives in full sun, in fertile,
Kurjenmiekka well-drained, neutral to slightly acidic soil with
French: Flambe, Iris, Iris Allemande, Iris pH 6–7. It needs light watering. It is widely
d’Allemagne, Iris Germanique grown as a garden ornamental in the temperate
German: Deutsche Schwertlilie, Echte regions of Australia and has become naturalised
Schwertlilie, Gelbe Schwertlilie, along roadsides, in waste areas and in bushland.
Himmelschwertel, Ritter-Schwertlilie, Türk
Hungarian: Kerti Nőszirom, Kék Nőszirom, Edible Plant Parts and Uses
Nepáli Nőszirom
India: Puskaramulam (Malayalam), Kombirei Orrisroot is often included as one of the many
(Manipuri), Haimavati, Mulam, Padma- ingredients of Ras el hanout, a blend of herbs
Pushkara, Parasikavaca, Puskaramulam and spices used across the Middle East and
(Sanskrit) North Africa, primarily associated with
Italian: Giaggiolo Paonazzo, Giaggiolo Moroccan cuisine (Surhone et al. 2011). Peeled
Maggiore rhizomes of I. germanica (orrisroots) can be
Japanese: Ayame, Hanashoubu, Kakitsubata used as a flavouring in ice cream, confectionery
Korean: Ailiseu Germanica and baked goods (Bender 2009). Orris is also an
Persian: Bikh-I-Banafshah ingredient in many brands of gin. Orrisroot has
Polish: Kosaciec Bródkowy, Kosaciec Niemiecki, been used in tinctures to flavour syrups; its taste
Kosaciec Ogrodowy is said to be indistinguishable from raspberry
Portuguese: Lírio-Cardano, Lírio-Cardeno, (Chase 1900). In France, starch of I. germanica
Lírio-Da-Alemanha, Lírio-Germânico, root had been recommended a famine food for
Lírio-Roxo extending bread flour, after removal of the bitter
Slovašcina: Bradata Perunika, Nemška Perunika, element (Parmentier 1781, cited by Freedman
Perunika Nemška 2009).
Slovencina: Kosatec Nemecký
Spanish: Iris, Lirio Cardeno, Lirio Común
Swedish: Trädgårdsiris Botany
Thai: M̀āntā Germanica
Tibetan: Su Dag Dkar Po Iris x germanica is a rhizomatous, perennial herb,
Turkish: Mor Süsen, Navruzu, Türk Süzeni growing to 120 cm high, forming a large clump
Vietnamese: Diên Vỹ German hoa tím to 30 cm wide (Plate 1). Rhizomes are homoge-
Welsh: Gellesgen Farfog neous, creeping on soil surface or to 10 cm depth,
usually many-branched, light brown, 1.2–2 cm in
diameter and smooth, with nodal rings; branches
may arise in the fan or as many as 15–20 nodes
Origin/Distribution are produced prior to active leaves. Stems are
green, 2–3-branched, solid, 60–120 cm × 1–1.5 cm
I. x germanica probably originated in central and glaucous. Leaves are purplish at base and
southern Europe and the Balkan Peninsula. Iris folded midrib to base, glaucous, ensiform and
germanica is considered to be a natural and fer- 45 cm long by 3.5 cm wide. Inflorescences with
Iris x germanica 29

Nutritive/Medicinal Properties

Rhizome Phytochemicals

The following compounds were isolated from I.

germanica rhizomes: irisolone (Dhar and Kalla
1972), 5,4′-dihydroxy-6,7-methylenedioxyiso-
flavone (irilone) (Dhar and Kalla 1973) and
2,4,6,4′-tetrahydroxybenzophenone (Dhar and
Kalla 1974). Acetovanillon (4-hydroxy-3-me-
thoxyacetophenon) and the known flavonoids
irisolidone, irigenin, irisolone, tectorigenin and
dihydroquercetin-7.3′-dimethylether were
detected in Iris germanica rhizomes (Pailer and
Franke 1973). Also, a few unknown compounds
were isolated and their structure elucidated as
1.3-dioxolo[4.5-g] [1]-benzopyran-8-on (=5.3′.
(III A); 9-methoxy-7-(3′.4′-dimethoxyphenyl)-
8H-1.3-dioxolo[4.5-g] [1]-benzopyran-8-on (=5.
III B); 9-hydroxy-7-(p-hydroxyphenyl)-8H-1.3-
dioxolo[4.5-g] [1]-benzopyran-8-on (=5.4′-dihy-
Plate 1 Orrisroot plant habit
droxy-6.7-methylenedioxyiosoflavone (IX); 5.7-
terminal unit are 2–3-flowered, and branch units methoxy-4H-1-benzopyran-4-on
1–2-flowered; spathes are green, sometimes with (=5.7.3′-trihydroxy-6.4′-dimethoxyisoflavone
purple base, and 2–5 cm, with narrow, scarious (XI B); 5.7-dihydroxy-3-(4′-hydroxy-3′-metho-
margins and tip. For the flowers, the perianth has xyphenyl)-6-methoxy-4H-1-benzopyran-4-on
shades of blue-violet, yellow, brown or white (=5.7.4′-trihydroxy-6.3′-dimethoxyisoflavone
(Plate 2) with various patterns of pigment distri- (XI C). The ethanol rhizome extract of I. german-
bution; floral tubes are 1–2.5 cm; sepals are ica afforded an isoflavone homotectoridin with
spreading, drooping or reflexed and have shades the structure 5, 4′-dihydroxy-8, 3′-dimethoxy-
of blue-violet, yellow, brown or white with pat- isoflavone-7-O-mono-D-glucoside and tectoridin
terned overlay of darker blue-violet, with white (Kawase et al. 1973). Acetovanillon, irigenin and
or yellow beard along midrib of claw and lower its glucoside iridin, β-sitosterol, α-amyrin and
part of limb; obovate limbs taper gradually to β-amyrin were isolated from the fresh rhizomes
claw, 6–7.5 by 4–5.5 cm; petals alternate with of I. germanica (El-Moghazzy et al. 1980).
sepals and are erect and obovate, 5–7 by Bicyclic triterpenoids, α-irigermanal and
4–5.5 cm, with short, 1.5–2 cm, channelled claw; γ-irigermanal, and a monocyclic triterpenoid, iri-
the ovary is bluntly trigonal, 1.5–2.5 cm and dogermanal, were the major extractable lipids
slightly wider than floral tube. Capsules are isolated from the rhizomes, constituting about
bluntly angled, 3-lobed borne on ends of stems 1 % of the fresh weight (Marner et al. 1982).
and branches and 3–5 × 2.5 cm containing From the rhizome, one new hexaoxygenated
reddish-brown, wrinkled, oval seeds, 3–4 mm in isoflavone, 5,3′-dihydroxy-4′,5′-dimethoxy-
2 rows per locule. 6,7-methylenedioxyisoflavone; two new polyox-
30 Iridaceae

Plate 2 White flowers

ygenated isoflavone glucosides, 5-hydroxy-6,4′- Three piscidal triterpenes named irisgermani-

dimethoxyisoflavone 7-glucoside and 5,3′- cals A, B and C and seven known iridal-type
dihydroxy-4′,5′-dimethoxyisoflavone 7-glucoside; tripenes, namely, iripallidal, iriflorental,
and the known isoflavonoids irisolidone, irigenin α-irigermanal, γ-irigermanal, isoiridogermanal,
and iridin and acetovanillone, sitosterol, α-amyrin 16-O-acetyl-isoirigermanal and α-dehydroiri-
and β-amyrin were isolated (Ali et al. 1983). germanal, were isolated (Ito et al. 1995). Iridal
Glucose content of I. germanica rhizomes, kept triterpenoids isolated from the rhizome included
under anoxia (absence of oxygen), rapidly and irisgermanicals A, B and C, isoiridogermanal,
dramatically decreased during the first 30 days and 16-O-acetylisoiridogermanal, a-irigermanal,
then increased at the same time as adenylate g-irigermanal, a-dehydroirigermanal, iridal, iri-
energy charge values started to decline (Hanhijarvi florental and iripallidal (Miyake et al. 1997).
1995). The amount of non-reducing sugars Four isoflavones glycoside iriskashmirianin
decreased gradually during the anoxic incubation. 4′-O-β-D-glucoside, nigricin 4′-O-β-D-glucoside,
Under aerobic control conditions, adenylate irilone 4′-O-β-D-glucoside and iridin were iso-
energy charge (AEC) of I. germanica rhizome tis- lated from the rhizomes (Atta-ur-Rahman et al.
sue was 0.81. Under anoxia the energy charge of I. 2002). A monocyclic triterpene ester iristecto-
germanica rhizome tissue remained above 0.6 for rone K was isolated from the rhizomes of Iris
4 days only. Large amounts of ethanol were found germanica from Turkey (Orhan et al. 2002). Six
in anoxic rhizome (0.06 M) after 8 days. Long- known isoflavones, irisolidone, irisolidone
lived free radicals were rapidly generated in I. ger- 7-O-α-D-glucoside, irigenin, irilone, iriflogenin
manica plant tissues when treated anoxically and and iriskashmirianin, were isolated from Iris ger-
subsequently returned to aerobic conditions manica rhizomes (Wollenweber et al. 2003). Five
(Crawford et al. 1994). The free radicals in I. ger- new di- and triglycosides, irigenin 7-[O-β-D-
manica rhizomes were extracted into aqueous glucopyranosyl-(1 → 6)-β-D-glucopyranoside];
alkali and found to be flavonoids including querce- 6-hydroxygenistein 4′-[O-β-D-glucopyranosyl-
tin, irisolone, selenone and derivatives of irigenin. (1 → 6)-β-D-glucopyranoside]; nigricin 4′-[O-β-
The main flavonoids had more reduced and/or D-glucopyanosyl-(1 → 6)-β-D-glucopyranoside];
more alkylated structures than those from I. ger- nigricin 4′-[O-β-D-glucopyanosyl-(1 → 2)-O-
manica grown under normal aerobic conditions. [α-L-rhamnopyranosyl-(1 → 6)]-β-D-
Semiquinone radical anions could readily be gen- glucopyranoside]; and 7-4′-{[2″-O-(4′′′′-acetyl-
erated from quercetin, and similar flavonoids with 2′′′′-methoxyphenyl)-β-D-glucopyranosyl]oxy}-
1,2-dihydroxy-, 1,4-dihydroxy- or trihydroxy-sub- 3′-(β-D-glucopyranosyloxy)phenyl]-9-methoxy-
stitution patterns, by interaction with superoxide. 8H-1,3-dioxolo[4,5-g]-[1 benzopyran-8-one-]
Iris x germanica 31

along with a known compound, nigricin 4′-(β-D- of isoflavone glucosides. Two novel compounds
glucopyranoside), were isolated from Iris ger- 5,2-dihydroxy-3-methoxy-6,7-
manica rhizomes (Atta-ur-Rahman et al. 2003a). methylenedioxyflavone and 5,7,2-trihydroxy-
Nine isoflavonoids: 5,7-dihydroxy-3-(3′-hydroxy 6-methoxyflavanone were isolated from the
-4′,5′dimethoxy)-8-methoxy-4H-1-benzopyran- fraction B of the rhizome chloroform extract
4-one; 5,7-dihydroxy-3-(3′-hydroxyl-4′, 5′-dim- (Singab et al. 2006). The methanol extract of Iris
ethoxy)-6-methoxy-4H-1-benzopyran-4-one; 5, germanica rhizome afforded one new compound,
7-dihydroxy-3-(4′-hydroxy)-6-methoxy-4H-1- 6,6-ditetradecyl-6,7-dihydrooxepin-2(3H)-one
benzopyrane-4-one; 5-hydroxy-3-(4′-hydroxy)- and five known compounds, 1-(2-(6′-hydroxy-2′-
6,7-methylenedioxy-4 H -1-benzopyran-4-one; methylcyclohex-1′-enyloxy)-5-methoxyphenyl)
5-hydroxy-3-(4′-methoxy)-6,7-methylenedioxy-4H- ethanone; 4-hydroxy-3-methoxyacetophenone;
1-benzopyran-4-one; 5-methoxy-3-(4′-hydroxy)- irisolone; irisolidone and 2-acetoxy-3,6-
6,7-methyenedioxy-4 H -1-benzopyran- 4-one; dimethoxy-1,4-benzoquinone (Asghar et al.
5,7-dihydroxy-3-(3′-hydroxy-4′-methoxy)-6- 2009).
methoxy-4H-1-benzopyran-4-one; 5,7-dihydroxy A total of 20 compounds identified as isofla-
-3-(3′-methoxy-4′-hydroxy)-6-methoxy-4H-1- vones, isoflavone glycosides and acetovanillone
benzopyran-4-one and isopeonol were isolated were isolated from the lipophilic and polar
from the rhizome (Atta-ur-Rahman et al. 2003b). extracts of Iris germanica rhizomes (Schütz et al.
Five new di- and triglycosides, irigenin 7-[O-β-D- 2011). A new isoflavone glycoside, iriflogenin-
glucopyranosyl-(1 → 6)- β - D - glucopyranoside] 4′-O-gentiobioside, was isolated. Nine isofla-
(1); 6-hydroxygenistein 4′-[O-β-D-glucopyranosyl- vones (totalling 180 mg/g) were identified in I.
(1 → 6)-β-D-glucopyranoside] (2); nigricin germanica resinoid: irigenin, iristectorigenin A,
4′-[O-β-D-glucopyanosyl-(1 → 6)-β-D- nigricin, nigricanin, irisflorentin, iriskumaonin
glucopyranoside] (3); nigricin 4′-[O-β-D- methyl ether, irilone, iriflogenin and irisolidone
glucopyanosyl-(1 → 2)-O-[α-L-rhamnopyranosyl- (Roger et al. 2012). The methanolic extract of
(1 → 6)]-β-D-glucopyranoside] (4) and 7-{4′- Iris germanica rhizomes yielded two new
{[2″-O-(4′′′′-acetyl-2′′′′-methoxyphenyl)-β-D- compounds, irigenin S and iriside A, together
glucopyranosyl]oxy}-3′-( β - D -glucopyran- with known compounds: stigmasterol, α-irone,
osyloxy)phenyl]-9-methoxy-8H-1, γ-irone, 3-hydroxy-5-methoxyacetophenone,
3-dioxolo[4,5-g]-[1 benzopyran-8-one-] (5), irilone, irisolidone, irigenin, stigmasterol-3-
along with a known compound, nigricin 4′-(β-D- O-β-D-glucopyranoside, irilone 4′-O-β-D-
glucopyranoside), were isolated from Iris ger- glucopyranoside and iridin (Ibrahim et al. 2012).
manica rhizomes (Nasim et al. 2003). Irigenin Two new compounds, 5-methoxy-3′,4′-
(5,7,3′-trihydroxy-6,4′,5′-trimethoxyisoflavone) dihydroxy-6,7-methylenedioxy-4H-1-
and iristectorigenin A (5,7,3′-trihydroxy-6,4′- benzopyran-4-one (iriskashmirianin A) and
dimethoxyisoflavone) along with their 7-O-β-D- 5,3′-dihydroxy-3-(4′-β-D-glucopyranosyl)-6,7-
glucosides, iridin and iristectorin A, respectively, methylenedioxy-4H-1-benzopyran-4-one (ger-
were found as the major components in adventi- manaism H), along with eight known compounds,
tious roots in the liquid medium, and the total namely, irilone, iriskumaonin methyl ether, iriflo-
isoflavone content was about 3.6 μmol per g fresh genin, irisolone, irifloside, irilone 4′-O-β-
weight in 3-week-old I. germanica cultures glucopyranoside, germanaism A and germanaism
(Akashi et al. 2005). The total isoflavone content B were isolated from the rhizomes (Xie et al.
(glucosides + aglycones) at 6 h after the start of 2013). New ceramides irisamides A and B and
CuCl2 treatment was 1.4-fold the initial value, isoflavone iridin S were isolated from the rhi-
and nearly the same content was maintained for zome (Mohamed et al. 2013). The methanol
48 h. Thus, the main effect of CuCl2 treatment extract of Iris germanica afforded a new benzene
appeared to be the induction of hydrolysis derivative 2′-methyl-6′-hydroxy cyclohexenyl-
32 Iridaceae

3-methyl-1-acetophenone ether and another (0.14 %); benzoic acid, 4-ethoxy-, ethylester
known benzene derivative isopenol together with (0.12 %); hexadecane (0.11 %); thiosulfuric acid,
two known isoflavones, irisolone and irisolidone S-(2aminoethyl) ester (0.1 %); tetradecane
(Asghar et al. 2010). Twenty-one compounds (0.08 %); and decane (0.06 %).
were isolated from the rhizomes of Iris german- Orrisroot is used as an aromatic agent. Iris
ica and identified as ombuin (1); 5, 3, 3′-trihy- pallida is the best for extractive purposes
droxy-7, 4′-dimethoxyflavanone (2); naringenin followed by I. germanica and I. florentina for
(3); cirsiliol-4′-glucoside (4); 3β, 4′-dihydroxy- botanical sources of orrisroot preparations (con-
7,3′-dimethoxyflavonone-5-O-β-D-- crete, liquid, oil, root extract) (Fenaroli 1994).
glucopyranoside (5); genistein (6); irilin D (7); Orris absolute consists mainly of isomeric irones
muningin (8); 5, 7, 4′-trihydroxy-6, 3′, 5′-trime- (Guenther 1952). Orris absolute (alcoholic
thoxyisoflavone (9); tectorigenin (10); irigenin extraction or distillation of orris concrete) was
(11); tectoridin (12); iridin (13); mangiferin (14); reported to have a yield of 0.03–0.05 % dried rhi-
irisxanthone (15); pyroglutamic acid (16); 2, 4′, zome and to contain 55–58 % ketones (irones),
6-trihydroxy-4-methoxybenzophenone-2-O-β-D- free or partially esterified fatty acids (methyl
glucoside (17); apocynin (18); androsin (19); myristate, methyl caprylate, methyl laurate,
β-sitosterol (20); and daucosterol (21) (Xie et al. methyl oleate, methyl linoleate), aldehydes
2014). (oleic, benzoic), alcohols (benzylic), terpene and
Eleven compounds were identified in the petro- sesquiterpene (Burdock 1994). Orris resinoid
leum ether extract (oil) of I. germanica: 9-hexa- (yield 1–3.3 %) was reported to contain 62–78 %
decanoic acid methyl ester; 9-octadecenoic acid ketones (irones) (Burdock 1994). Orris absolute
methyl ester; 8-octadecenoic acid methyl ester; prepared from Iris x germanica was reported to
11-octadecenoic acid methyl; 10-octadecenoic contain (Z)-a-irone (558.4–62.7 %), (Z)-γ-irone
acid methyl ester; 13-octadecenoic acid methyl (32.2–36.9), (E)-α-irone (1.7–3.9 %) and (Z)-β-
ester; 16-octadecenoic acid methyl ester; 1, irone (0.4–2.3 %) and orris essential oil to con-
2-benzenedicarboxylic acid diisooctyl ester; bis(2- tain (Z)-α-irone (57.6–66.2 %), (Z)-γ-irone
ethylhexyl) phthalate; methyl 6-methyl heptano- (33.8–39.4 %), (E)-α-irone (0.5–2.7 %) and (Z)-
ate; and nonanoic acid, 9-oxo-methyl ester (Asghar β-irone (0–2.2 %) (Galfre et al. 1993). (+)-cis-γ-
et al. 2011). Twenty-three compounds were identi- Irone and (+)-cis-α-irone were found to have the
fied in the essential oil of Syrian Iris germanica most interesting organoleptic properties. Orris
with myristic acid (61.42 %) as the major compo- essential oil is a cream-coloured solid and is
nent (Almaarri et al. 2013). The other sub-major often referred to as orris butter and often and
compounds obtained were elaidic acid methyl erroneously referred to as orris concrete which is
ester (9-octadecenoic acid methyl ester) (6.61 %), a dark, viscous material perhaps more correctly
lauric acid (5.69 %), octadecanoic acid methyl referred to as a resinoid.
ester (stearic acid methyl ester) (5.40 %), palmitic Irones extracted from Iris species are aromatic
acid methyl ester (5.10 %) and palmitic acid principles used in the perfume and flavour indus-
(4.87 %). The minor compounds were linoleic try to impart violet fragrance. Four major irone
acid methyl ester (1.19 %); linoleic acid (1.1 %); compounds (trans α-, cis α-, cis γ- and β-irones)
docosane (1.08 %);10-octadecenoic acid, methyl had been described (Naves et al. 1947; Schinz
ester (0.7 %); 3-methyl-1,2-benzoisoselenazple et al. 1947; Jaenick and Marner 1990) as respon-
(0.61 %); nonadecane (0.55 %); oleic acid sible for the characteristic scent of Iris oil. Natural
(0.53 %); sulphurous acid,2-propyl tetradecyl irones (terpenoids) in essential oil of Iris rhizomes
ester (0.28 %); α-irone (0.25 %); 2-propenoic were found to be formed by oxidative degrada-
acid, 3-4(-methoxy-phenyl)-2ethyl-hexyl ester tion of other higher terpenes C31-triterpenoids
(0.16 %); 1,2-benzenedicarboxylic acid, dioctyl produced by the plant (Krick et al. 1984).
ester (0.15 %); decanoic acid (capric acid) Triterpenoids called iridals were shown to be
Iris x germanica 33

irone precursors (Jaenick and Marner 1990). manica rhizomes (Bonfils and Sauvaire 1996).
Iridals originated from C30 olefin squalene Monocycloiridals were mostly represented in
(Jaenick and Marner 1990, Marner et al. 1988; young rhizomes, whereas cycloiridals appeared
Marner 1997). The incorporation of [2-14C]ace- to be the main iridal formed in the old ones. The
tate, [2-14C]mevalonate and [3H]squalene proved study of the ratio of free iridals (FI)/iridal esters
the squalenoid nature of the iridals (Marner et al. (IE) revealed a higher ratio in the youngest rhi-
1988). Methionine was readily incorporated into zomes. In addition 20 irone-related compounds
cycloiridals of Iris pallida dalmatica, thus indi- containing 10-16C-atoms were detected in orris
cating that the methylation of iridals via concrete of Moroccan origin (I. germanica)
S-adenosyl-L-methionine led to the formation of (Maurer et al. 1989). Iris germanica was found to
the irone moiety of the bicyclic compounds. The have a fresh mass of 1342 g/plant and a dry mass
cycloiridals, C31-triterpenoids, also served as of 437 g/plant and to contain 595 mg/kg DM
precursors of the irones (Marner et al. 1990). irone content that was composed of 79.1 % cis-γ-
Since 1982, more than 25 different iridal struc- irone and 20.9 % cis-α-irone (Firmin et al. 1998).
tures were characterised (Jannick and Marner An epoxidised iridal 18,19-epoxy-10-deoxyiridal
1990, Abe et al. 1991; Marner and Kerp 1992) was isolated from rhizomes of Iris germanica
and separated into 3 classes monocycloiridals, var. ‘Rococo’ (Bonfils et al. 1998). Ten enantio-
bicycloiridals and spiroiridals. Rhizomes of both pure isomers of irone were reported to be the
I. pallida varieties were found to contain 0.1– odoriferous principle of orrisroot oil, by means of
1.1 %, and I. germanica 0.1–0.5 % of iridals and enzyme-mediated approaches, starting from
their esters (Marner and Kerp 1992). Bonfils commercial racemic Irone alpha® (Brenna et al.
et al. (1996) found iridals were membrane 2003). A new iridal irigermanone and nine known
located. Two triterpenoids iripallidal and iriflo- congeners were isolated from the dried rhizomes
rental, known irone precursors, were found to be of German iris (Iris germanica) (Potterat et al.
solubilised by phosphatidylcholine. These cyclo- 2014). The compound was a structurally unique
iridals were found to be integrated within liposo- noriridal and possessed an unprecedented meth-
mal membranes and appeared to have a structural ylcarbonyl group instead of the α,β-unsaturated
role within cells comparable to that of sterols. aldehyde function typical for this group of
Iris germanica had high levels of (+)cis-α-irone triterpenes.
and low content of trans-α-irone, while I. pallida
had a high content of (-) cis-α-irone (Maurer
et al. 1989; Firmin et al. 1998). Dehydration of Other Plant Parts’ Phytochemicals
rhizome slices led to an increase of iridal esters
with a concomitant drop of free iridals, as well as A C-glycosyl flavone, embinin, is isolated from
phospholipids, in the drought-resistant species, the petals of I. germanica (Kawase and Yagishita
Iris germanica (Bonfils et al. 1994). Analysis of 1968). In Iris germanica, flower colour was
the intracellular membranes of Iris germanica found to be determined by two distinct biochemi-
rhizomes indicated high amounts of iridals when cal pathways, the anthocyanin biosynthetic and
compared to sterols and lower sterol amounts the carotenoid biosynthetics (Warburton et al.
than in other plant microsomes (Bonfils et al. 1978; Meckenstock 2005). In cyanic iris flowers,
1995). Incubation of a monocyclic iridal (a triter- the main pigments reported were delphinidin-
penoid from Iris species) with rat liver micro- type anthocyanins imparting blue, violet and
somes in the presence of NADPH led to the maroon colours, while in acyanic flowers, the
degradation of this substrate and to the genera- main pigments were carotenoids, producing
tion of 16-hydroxy-iridal (isoiridogermanal) orange, yellow and pink colours. A red iris does
(Bonfils et al. 1955). Iridal composition was also not naturally exist because of a deficiency in the
found to differ, depending on the age of Iris ger- synthesis of pelargonidin- and cyanidin-type
34 Iridaceae

anthocyanins, usually responsible for red and pounds was rated in the following order:
orange hues or that red carotenoids, such as lyco- BHA > BHT > α-tocopherol > > aqueous iris
pene, do not accumulate in large concentrations, extract > ethanol iris extract. The scavenging
thus producing only pink flowers, as opposed to effect of the aqueous and ethanol iris extracts and
red ones (Jeknić et al. 2014). With a goal of the reference compounds on the DPPH* radical
developing iris cultivars with red flowers, they decreased in the following order:
transformed a pink iris I. germanica, ‘Fire Bride’, BHA > α-tocopherol > BHT > aqueous
with a bacterial phytoene synthase gene (crtB) iris > ethanol iris extract. These results indicated
from Pantoea agglomerans under the control of that both iris extracts produced a noticeable scav-
the promoter region of a gene for capsanthin– enging of free radicals; free radical scavenging
capsorubin synthase from Lilium lancifolium activity also increased with the concentration of
(Llccs). They showed that ectopic expression of a iris extracts. The metal chelating effect of both
crtB could be used to successfully alter the colour iris extracts and the reference compounds
of certain flower parts in I. germanica ‘Fire decreased in the following order: aqueous iris
Bride’ and produce new flower traits. The colour extract > ethanol iris extract > BHA > α-tocophero
changes were confirmed primarily due to the l > BHT. The results indicated that iris had in-
accumulation of lycopene. Other carotenoids and vitro antioxidant properties, which could be the
chlorophyll found in both control and crtB peri- major factor responsible for the inhibition of lipid
gon (standards and falls), ovaries and flower stalk peroxidation. The petroleum ether extract (oil) of
were neoxanthin, violaxanthin, lutein, lycopene, I. germanica exhibited good DPPH scavenging
chlorophyll a, chlorophyll b and α + β-carotene. activity but showed moderate inhibition in lin-
Carotenoids β-carotene 7.1 % (4.3 mg), lutein oleic acid oxidation as compared to BHT (Asghar
87.8 % (53.2 mg), neochrome (zeaxanthin furan- et al. 2011). The radical scavenging activity of
oxide) 3.5 % (2.1 mg) and 7 minor constituents the ethanolic extracts of the aerial parts and rhi-
1.6 % (1 mg) were found in the green aerial parts zomes of I. germanica were found to have IC50
of I. germanica (Bucheckerand and Liaaen- values of 5.38 and 12.3 mg/ml, respectively,
Jensen 1975). From the leaves of I. pallida, 0.1– whereas total antioxidant activity of the extracts
0.7 % of iridals and their esters were extracted (at 3.15 mg/ml) was 98.7 % and 97.4 %, respec-
and from I. germanica leaves 0.05–0.3 % (Marner tively, and total phenolic content was 267.36 and
and Kerp 1992). 331.96 8 mg gallic acid equivalent/g extract,
respectively (Burcu et al. 2014). HPLC analysis
of phenolic compounds identified protocatechuic
Antioxidant Activity acid (0.356 mg/g extract), chlorogenic acid
(0.164 mg/g extract) and ferulic acid (0.164 mg/g
Both aqueous and ethanol I. germanica rhizome extract) as the main phenolic acids contained in
extracts exhibited strong total antioxidant activ- the extract of the aerial parts of I. germanica,
ity, showing 95.9, 88.4, 79.9 % and 90.5, 78.0, whereas chlorogenic acid (2.44 mg/g extract),
65.3 % inhibition on peroxidation of linoleic acid (+)-catechin (2.14 mg/g extract) and ferulic acid
emulsion at concentrations of 10, 30 and 50 μg/ (0.452 8 mg/g extract) were identified as the main
ml, respectively, and in all cases exceeded the phenolic acids contained in the extract of the
effect of 30 μg/ml α-tocopherol solution (36.6 %) rhizome.
(Nadaroğlu et al. 2007). Both extracts also pos-
sessed effective reducing power and exhibited
free radical scavenging, superoxide anion radical Anticancer Activity
scavenging, hydrogen peroxide scavenging and
metal chelating activities at concentrations of 20, Isoflavones from I. germanica rhizomes, irigenin,
40 and 60 μg/ml. The reducing power of aqueous irilone and iriskashmirianin exhibited moderate
and ethanol iris extracts and reference com- activity as inducers of NAD(P)H:quinone reduc-
Iris x germanica 35

tase (QR) in cultured mouse Hepa 1c1c7 cells, (EAC) cancer cell line with IC50 of 20.9 7 and
with CD values (concentration required to double 4.3 μM for 3-(4, 5-dimethylthiazole-2-yl)-2,
the specific activity of QR) of 3.5–16.7 μM, 5-diphenyltetrazoli-umbromide (MTT) and ATP
whereas weak activity was observed with com- assay methods, respectively (Xie et al. 2013).
pounds iriflogenin and iriskashmirianin in the
radical (DPPH) scavenging bioassay (IC50 values
89.6 and 120.3 μM, respectively) (Ito et al. 1995). Antimutagenic Activity
Iridals extracted from Iris germanica exhibited
antitumour effects in-vitro against human tumour The ethanolic extract of the aerial parts of I. ger-
cell lines: A2780 and K562 (and for each one, a manica exhibited antimutagenic effects at 3 and
drug-sensitive and a drug-resistant cell line) with 0.3 mg/plate concentrations as assessed using the
IC50 values of 0.1–5.3 μg/ml (Bonfils et al. 2001). Ames Salmonella typhimurium/microsome
Some of them were shown to be more effective mutagenicity test (Burcu et al. 2014). The rhi-
than doxorubicin. Studies by Halpert et al. (2011) zomes also exhibited antimutagenic effects at
found that the lipidic extract from Iris germanica 1.5, 0.15 and 0.015 mg/plate concentrations.
was able to increase HeLa cell area and adhesion
and augment the formation of actin stress fibres.
This effect was not observed when Ref52 fibro- Antimicrobial Activity
blasts were tested and was not the result of dis-
ruption of microtubules. Further, the increase in The chloroform and ethyl acetate extracts of I.
cell area was Rac1 dependent, and the iris extract germanica rhizomes exhibited bactericidal
led to slight Rac activation. The increase in HeLa activity, while the petroleum ether extract did
cell area in the presence of iris extract was not exhibit any bactericidal, fungicidal and
accompanied by impairment of cell migration insecticidal activities (Orhan et al. 2003 ). It
and arrest of the cell cycle at G1 although there is was also inactive in the brine shrimp toxicity
involvement of Rac1. The active compounds test, whereas it showed significant phytotox-
were found to iridals, a known group of triterpe- icity against the plant Lemna aequinoctialis.
noid. Purified iripallidal was able to increase cell The hexane fraction of the methanol rhizome
area of both HeLa and SW480 cells. extract showed significant inhibitory activity
With respect to antitumour-promoting poten- against Fusarium solani (70 %) and moderate
tial based on anti-inflammatory mechanisms, activity against Trichophyton longifusus
none of the compounds demonstrated significant (50 %) and Microsporum canis (30 %), while
activity in the concentration range tested. Iris the ethyl acetate and chloroform fractions
germanica ceramides irisamides A and B were exhibited moderate activity against the tested
active against mouse lymphoma L5178Y and fungi (Asghar et al. 2009 ). It was observed
cervical cancer HeLa cell lines, but inactive that the growth of bacteria (Pasteurella multo-
against PC12 rat pheochromocytoma (Mohamed cida, Escherichia coli , Bacillus subtilis,
et al. 2013). Irisamides A and B gave 76 % and Staphylococcus aureus) and fungi ( Ganoderma
63 % growth suppression against the L5178Y lucidum, Aspergillus flavus, A. niger,
and HeLa cell lines at a concentration of Alternaria alternata) were inhibited in I. ger-
12.67 mM as well as 68 % and 49 % at a concen- manic a petroleum ether extract (oil) as com-
tration of 3.81 mM, with ED50(s) of 0.91 and pared to the reference standards (rifampicin/
2.40 mM, respectively. The isoflavone iridin S terbinafine) (Asghar et al. 2011). The metha-
was inactive against the three cell lines. Among nolic rhizome extract showed antimicrobial
the compounds isolated from the rhizome, the activity against Staphylococcus aureus,
isoflavone iriskashmirianin A possessed the best Serratia marcescens, Escherichia coli,
cytotoxic activity in Ehrlich’s ascites carcinoma Candida albicans and Aspergillus flavus with
36 Iridaceae

the highest antimicrobial effect against S. IL-2, IFN-gamma and Th2: IL-4 and IL-5 in a
aureus (Ibrahim et al. 2012 ). dose-dependent manner, as studied by the flow
cytometric method. Oral administration of the
isoflavones in Balb/c mice at doses of 0.1–0.8 mg/
Antihyperlipidemic Activity kg per oral dose showed irisolidone to possess
stimulatory activity on T lymphocytes (CD4+
Administration of the ethanolic extract of Iris and CD8+ cells) and Th1: IL-2, IFN-gamma
germanica rhizomes for 10 weeks significantly cytokine production, while irilone acted as an
lowered the lipid components especially the cho- immunosuppressant for T lymphocytes (CD4+
lesterol and triglycerides in rats fed a high-fat and CD8+ cells) and T cell cytokines, namely,
diet (Choudhary et al. 2005) Th1: IL-2, IFN-gamma and Th2: IL-4 and IL-5.
The methylated products of both isoflavones
showed a similar trend to that of their parent
Drug Metabolising Activity compounds, but their activity was drastically
decreased revealing the importance of free phe-
Six isoflavones irisolidone, irisolidone 7-O-α-D- nolic groups for their immunomodulating
glucoside, irigenin, irilone, iriflogenin and iris- activities.
kashmirianin isolated from Iris germanica
rhizomes were shown to be potent inhibitors of
cytochrome P450 1A activity with IC50 values in Antimalarial Activity
the range of 0.25–4.9 μM (Wollenweber et al.
2003). Iridal, a triterpenoidic compound isolated from I.
germanica, exhibited antiplasmodial activity
(Benoit-Vical et al. 2003). The IC50 obtained in-
Anti-inflammatory Activity vitro on human malaria Plasmodium falciparum
chloroquine-resistant and chloroquine-sensitive
Isoflavonoids isolated from I. germanica rhizome strains ranged from 1.8 to 26.0 μg/ml and the
exhibited ant-inflammatory activity determined ED50 in-vivo for P. vincke was about 85 mg/kg/
by a spectrophotometric assay using the activated day by intraperitoneal route. Iridal presented an
human neutrophils (Attar-ur-Rahman et al. antiplasmodial activity similar to that obtained
2003b). The methanolic rhizome extract and the with extracts from the plant Azadirachta indica
isolated flavonoids exhibited potent anti- classically taken as reference in malaria
inflammatory effects by suppressing hind paw phytomedicine
oedema (skin oedema) induced by 4 % formalin
in albino rats (Ibrahim et al. 2012). Irigenin S
showed activity similar to that of Molluscicidal Activity
Iris germanica rhizome chloroform extract
showed the highest molluscicidal activity
Immunomodulatory Activity (LC90 = 1.26 mg/l) against Biomphalaria alexan-
drina snails among the tested extracts of the rhi-
Two isoflavones, 5,7-dihydroxy-6,4′-dimetho- zomes (Singab et al. 2006). Fraction B prepared
xyisoflavone (irisolidone) and 5,4′-dihydroxy- from the chloroform extract was the most potent
6,7-methylenedioxyisoflavone (irilone) isolated molluscicide (LC90 = 0.96 mg/l); in addition, it
from Iris germanica, exhibited immunomodula- showed a significant heart rate reduction in the
tory activities (Nazir et al. 2009). They influenced snail after a 6- to 24-h exposure period. It also
the production of T lymphocytes (CD4+ and displayed a significant level of cercaricidal poten-
CD8+ cells) and T cell cytokines, namely, Th1: tial in a time-concentration-dependent manner.
Iris x germanica 37

Piscidal Activity a powerful cathartic and used for the treatment of

dropsy (Grieve 1971). Orrisroot is also employed
Among the iridals isolated from the rhizome, the for complaints of the lungs, coughs and hoarse-
most potent piscidal activity was observed for iri- ness, bronchitis and chronic diarrhoea. If taken in
florental with medial tolerance (TLm) value after large doses, it can cause nausea, vomiting, purg-
24 h of 0.1 μg/ml (Ito et al. 1995). The TLm of the ing and colic (Grieve 1971).
bicyclic iridals irisgermanical A was 0.8 μg/ml
and irisgermanicals Band C were 3 μg/ml.
The monocyclic iridals, isoiridogermanal and Other Uses
16-O-acetyl-isoirigermanal, were inactive. Of the
iridals isolated from I. germanica, iriflorental, iri- Iris resinoid obtained from Iris germanica or Iris
pallidal and γ-irigermanal exhibited a potent pallida rhizomes is widely used in the perfume
piscidal activity (using killifish, Oryzias latipes) industry (Roger et al. 2012). Whole roots of
at a concentration of less than 1 μg/mL (Miyake Orris, resembling the human form, are used in
et al. 1997). voodoo performances, and the powdered root is
an ingredient in ‘love potions’. A black dye is
obtained from the roots and a blue dye from the
Allergy Issue flowers (Grae 1974). The seeds are used as rosary
beads (Usher 1974). Powdered orrisroot is some-
Allergic manifestations caused by the use of a times put into rinsing water in laundries and
dentifrice containing orrisroot (Iris × germanica imparts a refreshing and fragrant scent to the
L. nothovar. Florentina) powder was reported by linen (Grieve 1971). Orrisroot, mixed with anise,
Winter (1948). was used in England as a perfume for linen as
early as 1480. The root is a source of orris pow-
der which is much used as a fixative and base
Toxicological Studies note in perfumery and pot-pourri, as an ingredi-
ent of dentifrices, toothpastes, breath fresheners,
The acute oral LD50 value for orris absolute in face powders, foot powders, sachet powder,
rats was reported as 9.4 g/kg (Moreno 1972). violet-scented soaps and cachous and as a food
Undiluted orris absolute applied to the backs of flavouring. Orris oil is used commercially in the
hairless mice and swine was found to be not irri- preparation of the finest scents and is also blended
tating (Urbach and Forbes 1972). No phototoxic with artificial violet perfumes, the odour of which
effects were reported for undiluted orris absolute it renders more subtle (Grieve 1971).
on hairless mice and swine. A patch test using
orris at full strength for 24 h produced no irrita-
tion reactions in 20 subjects (Katz 1946). Orris Comments
absolute tested at 3 % in petrolatum produced no
irritation after a 48-h closed-patch test on human Bearded iris is propagated by division or cutting
subjects (Kligman 1972). Orris produced no pri- from the original rhizome and by seed.
mary irritation in a repeated insult patch test on
43 human subjects (Majors 1972).
Selected References

Traditional Medicinal Uses Abe F, Chem RF, Yamauchi T (1991) Iridals from
Belamcanda chinensis and Iris japonica.
Phytochemistry 30:3379–3382
The root is diuretic, emetic, expectorant and Akashi T, Ishizaki M, Aoki T, Ayabe SI (2005) Isoflavonoid
mildly purgative (Grieve 1971; Usher 1974; production by adventitious-root cultures of Iris ger-
Launert 1981; Chiej 1984). Juice from the root is manica (Iridaceae). Plant Biotechnol 22(3):207–215
38 Iridaceae

Ali AA, El-Emary NA, El-Moghazi MA, Darwish FM, Brenna E, Claudio Fuganti C, Stefano Serra S (2003)
Frahm AW (1983) Three isoflavonoids from Iris ger- From commercial racemic fragrances to odour active
manica. Phytochemistry 22(9):2061–2063 enantiopure compounds: the ten isomers of irone.
Almaarri K, Zedan TA, Albatal N (2013) Chemical analy- Comp Rend Chim 6(5–6):529–546
sis of essential oils of some Syrian wild Iris species. Buchecker R, Liaaen-Jensen S (1975) The carotenoid pat-
Am J Biochem Mol Biol 3:38–49 tern in Iris germanica. Phytochemistry 14(3):851–852
Asghar SF, Aziz S, Habib-ur-Rehman, Ahmed I, Hussain Burcu B, Aysel U, Nurdan S (2014) Antimicrobial, anti-
H, Atta-ur-Rahman, Choudhary MI (2009) Secondary oxidant, antimutagenic activities, and phenolic com-
metabolites isolated from Iris germanica. Rec Nat pounds of Iris germanica. Ind Crop Prod 61:526–530
Prod 3(3):139–152 Burdock GA (1994) Fenaroli’s handbook of flavor ingre-
Asghar SF, Habib-ur-Rehman, Atta-ur-Rahman, dients. 3rd ed, vol 1. CRC Press, Boca Baton
Choudhary MI (2010) Phytochemical investigations Chase AW (1900) Dr. Chase’s recipes; or, information for
on Iris germanica. Nat Prod Res 24(2):131–139 everybody: an invaluable collection of about eight
Asghar SF, Habib-ur-Rehman, Choudahry MI, Atta-ur- hundred practical recipes, 40th edn. R. A. Beal
Rehman (2011) Gas chromatography-mass spectrom- Publisher, Ann Arbor. 648 pp
etry (GC-MS) analysis of petroleum ether extract (oil) Chiej R (1984) The Macdonald encyclopaedia of medici-
and bio-assays of crude extract of Iris germanica. Int nal plants. Macdonald & Co, London, 447 pp
J Genet Mol Biol 3(7):95–100 Choudhary MI, Naheed S, Jalil S, Alam JM (2005) Effects
Atta-ur-Rahman, Nasim S, Baig I, Jahan IA, Sener B, of ethanolic extract of Iris germanica on lipid profile
Orhan I, Choudhary MI (2002) Isoflavonoid glyco- of rats fed on a high-fat diet. T Ethnopharmacol
sides from the rhizomes of Iris germanica. Chem 98(1):217–220
Pharm Bull 50(8):1100–1102 Crawford MMR, Lindsay DA, Walton JC, Wollenweber-
Atta-ur-Rahman, Nasim S, Baig I, Orhan I, Sener B, Ratzer B (1994) Towards the characterization of radi-
Ayanoglu F, Choudhary MI (2003a) Isoflavonoid gly- cals formed in rhizomes of Iris germanica.
cosides from the rhizomes of Iris germanica. Helv Phytochemistry 37(4):979–985
Chim Acta 86:3354–3362 Dhar KL, Kalla AK (1972) Isoflavones of Iris kumaonensis
Atta-ur-Rahman, Nasim S, Baig I, Jalil S, Orhan I, Sener and I. germanica. Phytochemistry 11(10):3097–3098
B, Choudhary MI (2003b) Anti-inflammatory isofla- Dhar KL, Kalla AK (1973) A new isoflavone from Iris
vonoids from the rhizomes of Iris germanica. germanica. Phytochemistry 12(3):734–735
J Ethnopharmacol 86(2):177–180 Dhar KL, Kalla AK (1974) 2,4,6,4′-Tetrahydroxy-
Bender DA (2009) A dictionary of food and nutrition, 3rd benzophenone in Iris germanica. Phytochemistry
edn. Oxford University Press, Oxford 13(12):2894
Benoit-Vical F, Imbert C, Bonfils JP, Sauvaire Y (2003) El-Moghazzy AM, Ali AA, El-Emary NA, Darwish FM
Antiplasmodial and antifungal activities of iridal, a (1980) Isoflavones from the rhizomes of Iris german-
plant triterpenoid. Phytochemistry 62(5):747–751 ica. Fitoterapia 51(5):237–239
Bonfils JP, Sauvaire Y (1996) Localization of iridals in Firmin L, Courtois D, Pétiard V, Ehret C, Lerch K (1998)
Iris germanica rhizomes. Phytochemistry Evaluation of the natural variability in irone content
41:1281–1285 and selection of Iris sp. for perfume production. Hortic
Bonfils JP, Bonfils C, Larroque C, Baccou JC, Sauvaire Y Sci 133(6):1046–1047
(1955) 16-hydroxylation of iridal (triterpenoid from Freedman RL (2009) Famine foods database. Purdue
Iris species) mediated by cytochrome P450. Nat Prod University, West Lafayette, http://www.hort.purdue.
Lett 6(1):15–22 edu/newcrop/FamineFoods/ff_home.html
Bonfils JP, Sauvaire Y, Baissac Y, Marner FJ (1994) Iridal Galfre A, Martin P, Petrzilka M (1993) Direct enantiose-
levels in Iris rhizomes—effects of wounding and lective separation and olfactory evaluation of all irone
dehydration. Phytochemistry 37(3):701–705 isomers. J Essent Oil Res 5:265–277
Bonfils JP, Bonfils C, Larroque C, Surjus A, Gleize D, Grae I (1974) Nature’s colors – dyes from plants.
Sauvaire Y (1995) Lipid composition of microsomes MacMillan Publishing Co, New York
of Iris germanica rhizomes. Phytochemistry Grieve M (1971) A modern herbal. Penguin, 2 vols. Dover
38(3):585–587 Publications, New York, 919 pp
Bonfils JP, Sauvaire Y, Maurin L (1996) Evidence of Guenther E (1952) Oil of orris root. In: Nostrand V (ed)
cycloiridals as membrane constituents: effects on flu- The essential oils, vol 4. Reinhold, New York,
idity patterns compared to those of cholesterol. Planta pp 79–103
200(3):353–357 Halpert M, Abu-Abied M, Avisar D, Moskovitz Y,
Bonfils JP, Franz-Josef Marner FJ, Yves Sauvaire Y Altshuler O, Cohen A, Weissberg M, Riov J, Gottlieb
(1998) An epoxidized iridal from Iris germanica var HE, Perl A, Sadot E (2011) Rac-dependent doubling
“Rococo”. Phytochemistry 48(4):751–753 of HeLa cell area and impairment of cell migration
Bonfils JP, Pinguet F, Culine S, Sauvaire Y (2001) and cell cycle by compounds from Iris germanica.
Cytotoxicity of iridals, triterpenoids from Iris, on Protoplasma 248(4):785–797
human tumor cell lines A2780 and K562. Planta Med Hanhijarvi AM (1995) Comparison of carbohydrate utili-
67(1):79–81 zation and energy charge in the yellow flag iris (Iris
Iris x germanica 39

pseudacorus) and garden iris (Iris germanica) under on modified cyclodextrins. Helv Chim Acta
anoxia. Physiol Plant 939:493–497 73:2165–2170
Henderson NC (1992) What is Iris germanica? Bull Am Maurer B, Hauser A, Froidevaux J-C (1989) New irone-
Iris Soc 286:6–11 related constituents from the essential oil of Iris ger-
Henderson NC (2002) Iris Linnaeus. In: Flora of North manica L. Helv Chim Acta 72:1400–1415
America Editorial Committee (eds) 1993+. Flora of Meckenstock DH (2005) Breeding red irises: the carot-
North America North of Mexico, vol 26. New York/ enoids., Collierville. 163 pp
Oxford, pp 371–376 Miyake Y, Ito H, Yoshida T (1997) Identification of iridals
Ibrahim SR, Mohamed GA, Al-Musayeib NM (2012) as piscicidal components of Iridaceous plants and their
New constituents from the rhizomes of Egyptian Iris conformations associated with CD spectra. Can
germanica L. Molecules 17(3):2587–2598 J Chem 75(6):734–741
Ito H, Miyake Y, Yoshida T (1995) New piscicidal triter- Mohamed GA, Ibrahim SR, Ross SA (2013) New cerami-
penes from Iris germanica. Chem Pharm Bull des and isoflavone from the Egyptian Iris germanica
43(7):1260–1262 L. rhizomes. Phytochem Lett 6(3):340–344
Jaenicke L, Marner FJ (1990) The irones and their origin. Moreno OM (1972) Report to RIFM (Research Institute
Pure Appl Chem 62:1365–1368 for Fragrance Materials) May 4, 1972; cited by
Jeknić Z, Jeknić S, Jevremović S, Subotić A, Chen TH Opdyke DLJ (ed) op.cit
(2014) Alteration of flower color in Iris germanica L. Nadaroğlu H, Demir Y, Demir N (2007) Antioxidant and
‘Fire Bride’ through ectopic expression of phytoene radical scavenging properties of Iris germanica.
synthase gene (crtB) from Pantoea agglomerans. Pharm Chem J 41:409–415
Plant Cell Rep 33:1307–1321 Nasim S, Baig I, Orhan I, Sener B, Ayanoglu F, Choudhary
Katz AE (1946) Dermal irritating properties of essential MI (2003) Isoflavonoid glycosides from the rhizomes
oils and aromatic chemicals. Spice Mill 69:46–48 of Iris germanica. Helv Chim Acta
Kawase A, Yagishita K (1968) On the structure of a new 86(10):3354–3362
c-glycosyl flavone embinin, isolated from the petals of Naves Y-R, Grampoloff AV, Bachmann P (1947) Etudes
Iris germanica Linnaeus. Agric Biol Chem sur les matières végétales volatiles L. Etudes dans les
32(4):537–538 séries des méthyl-3-linalols, des méthyl-3-citrals et
Kawase A, Ohta N, Yagishita K (1973) On the chemical des méthyl-6-ionones. Helv Chim Acta 30:1599–
structure of a new isoflavone glucoside, homotectori- 1613. (In French)
din, isolated together with tectoridin from the rhi- Nazir N, Koul S, Qurishi MA, Taneja SC, Ahmad SF,
zomes of Iris germanica Linnaeus. Agric Biol Chem Khan B, Bani S, Qazi GN (2009) Immunomodulatory
37(1):145–150 activity of isoflavones isolated from Iris germanica
Kligman AM (1972) Report to RIFM (Research Institute (Iridaceae) on T-lymphocytes and cytokines. Phytother
for Fragrance Materials), Feb. 18, 1972; cited by Res 23(3):428–433
Opdyke DLJ (ed) (1979) Monographs on fragrance Orhan I, Şener B, Hashimoto T, Asakawa Y, Özgüven M,
raw materials. Oxford/New York, 732 pp. cited by Ayanoğlu F (2002) Ristectorone K, a novel monocy-
Opdyke DLJ (ed) (1979) Monographs on fragrance clic triterpene ester from Iris germanica rhizomes
raw materials. Oxford/New York, 732 pp growing in Turkey. Fitoterapia 73(4):316–319
Krick W, Marner FJ, Janicke L (1984) On the Orhan I, Nasim S, Tener B, Ayanoglu F, Özgüven M,
Stereochemistry of natural irones, dihydroirones, and Choudhary MI (2003) Two isoflavones and bioactivity
their precursors. Helv Chim Acta 67:318–324 spectrum of the crude extracts of Iris germanica rhi-
Launert E (1981) Edible and medicinal plants. Hamlyn, zomes. Phytother Res 17(5):575–577
London Pailer M, Franke F (1973) Űber inhaltstoffe der Iris ger-
Majors PA (1972) Report to RIFM, March 21, 1972; cited manica (Schwertlilie). Monatsh Chem 104:1394–
by Opdyke DLJ (ed) op.cit 1408 (In German)
Marner FJ (1997) Iridals and cycloiridals, products of an Parmentier A (1781) Recherches sur les végétaux nourri-
unusual squalene metabolism in sword lilies sans qui, dans les temps de disette, peuvent remplacer
(Iridaceae). Curr Org Chem 1:153–186 les alimens ordinaires. Avec de nouvelles observations
Marner FJ, Kerp B (1992) Composition of iridals, unusual su la culture des pommes de terre. Paris, France:
triterpenoids from sword-lilies, and the seasonal Imprimerie Royale, 599 pp. cited by Freedman R
dependence of their content in various parts of differ- (2009) Famine Foods.
ent Iris species. Z Naturforsch C 47(1-2):21–25 newcrop/faminefoods/ff_families/aristolochiaceae.
Marner FJ, Krick W, Gellrich B, Jaenicke L, Winter W html
(1982) Irigermanal and iridogermanal. Two new triter- Potterat O, Herzog C, Raith M, Ebrahimi SN, Hamburger
penoids from rhizomes of Iris germanica L. J Org M (2014) Irigermanone, a noriridal with unprece-
Chem 47:2531–2538 dented methylketone function, from Iris germanica.
Marner FJ, Gladtke D, Jaenicke L (1988) Studies on the Helv Chim Acta 97(1):32–38
biosynthesis of iridals and cycloiridals. Helv Chim Roger B, Jeannot V, Fernandez X, Cerantola S, Chahboun
Acta 71:1331–1338 J (2012) Characterisation and quantification of flavo-
Marner FJ, Runge T, König WA (1990) Separation of noids in Iris germanica L. and Iris pallida Lam. resin-
enantiomeric irones by gas-liquid chromatography oids from Morocco. Phytochem Anal 23(5):450–455
40 Iridaceae

Salt Lake City Winter GR (1948) Allergic manifestations Urbach F, Forbes PD (1972) Report to RIFM, May 26,
caused by the use of a dentifrice containing orris root 1972; cited by Opdyke DLJ (ed) op.cit
powder. J Periodontol 19(3):108 Usher G (1974) A dictionary of plants used by man.
Schinz H, Ruzicka L, Seidel CF, Tavel C (1947) Produits Constable, London. 619 pp
à odeur de violette. Synthèse de la d, l-α-irone. Helv Warburton B, Warburton BA, Hamblen M (eds) (1978)
Chim Acta 30:1810–1814. (In French) The world of irises. The American Iris Society,
Schütz C, Quitschau M, Hamburger M, Potterat O (2011) New York. 494 pp
Profiling of isoflavonoids in Iris germanica rhizome Wollenweber E, Stevens JF, Klimo K, Knauft J, Frank N,
extracts by microprobe NMR and HPLC-PDA-MS Gerhäuser C (2003) Cancer chemopreventive in vitro
analysis. Fitoterapia 82(7):1021–1026 activities of isoflavones isolated from Iris germanica.
Singab ANB, Ahmed AH, Sinkkonen J, Ovcharenko V, Planta Med 69(1):15–20
Pihlaja K (2006) Molluscicidal activity and new flavo- Xie GY, Qin XY, Liu R, Wang Q, Lin BB, Wang GK, Xu
noids from Egyptian Iris germanica L. (var. alba). Z GK, Wen R, Qin MJ (2013) New isoflavones with
Naturforsch C 61(1–2):57–63 cytotoxic activity from the rhizomes of Iris germanica
Surhone LM, Tennoe MT, Henssonow SF (2011) Orris L. Nat Prod Res 27(23):2173–2177
root. Betascript Publishing, Beau Basin. 100 pp Xie GY, Chen YJ, Wen R, Xu JY, Wu SS, Qin MJ (2014)
The Plant List (2013) Version 1.1. Published on the Chemical constituents from rhizomes of Iris german-
Internet; Accessed 10 ica. Zhongguo Zhong Yao Za Zhi 39(5):846–850. (In
Aug 2014 Chinese)
Stachys affinis

Scientific Name Bulgaria: Ranilist

Chinese: Gan Lu Zi, Kan-Lu-Tzu Cao-Shi-Can
Stachys affinis Bunge (Sweet As Dew), T’sao-Shih-Ts’an
(Herbaceous Rock Silk Worm), Di-Gu-Niu,
Ti-Ku-Niu (Ground Slug), Bao-Ta-Cai, Pao-
Synonyms T’a-Ts’ai (Precious Pagoda Vegetable), Kon
Loh (Cantonese)
Stachys geobombycis var. alba C.Y.Wu & Czech: Čistec Hlíznatý
H.W.Li, Stachys pauciflora Benth., Stachys Danish: Jcrosne, Apankartofler, Kinaskok,
sieboldii Miquel, Stachys sieboldii var. alba Knold-Galtetand, Stachysknolde
(C.Y.Wu & H.W.Li) H.B. Chen, Stachys sieboldii Dutch: Chinese Artisjok, Crosne Wortelknolletjes,
var. glabrescens C.Y.Wu & H.W.Li, Stachys Japanse Andoorn
sieboldii var. malacotricha Hand.-Mazz., Stachys Eastonian: Mugul-Nõianõges
tuberifera Naudin Finland: Mukulapähkämö
French: Crosnes, Crosne Du Japon, Épiaire
Affine, Stachys Tubereux
Family German: Chinesische Artischocke, Crosnes De
Japon, Japanische Kartoffel, Japanknollen,
Lamiaceae Knollen-Ziest, Lippenblütler, Stachys,
Greek: Stocchis
Common/English Names Hebrew: Eshbal
Italian: Betonica Bianca, Stregona Giapponese,
Artichoke Betony, Chinese Artichoke, Crosne, Tuberina
Japanese Artichoke, Knotroot Japanese: Chorogi
Korean: Mongulsuso
Lithuanian: Gumbuotoji Notra
Vernacular Names Macedonian: Čistec, Japonska Artičoka;
Malaysia: Tung-Tung Chow (Cantonese)
Mongolian: Argui
Afrikaans: Sjinese Artisjok Norwegian: Knollsvinerot, Svinerot
Brazil: Estachida Do Japão (Portuguese) Polish: Czysciec Bulwiasty

© Springer International Publishing Switzerland 2016 41

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_3
42 Lamiaceae

Portuguese: Alcachofra Chinesa, Crosne Do Edible Plant Parts and Uses

Japao, Estaque, Rab-De-Raposa, Urtiga
Japonesa The edible tuber is used as vegetable (Facciola
Slovašcina: Sieboldov Čišljak 1990; Larkcom 1991; van den Burgh 1996; Hu
Spanish: Alcachofa China, Alachofra Tuberose, 2005). Chinese artichokes are valued as a food in
Crosne Del Japón, Estaquídea Del Japón, the Chinese Five Systems, for feeding and
Ortigs Japonesa strengthening the respiratory and lymphatic sys-
Swedish: Korogi tem. They are an attractive, gourmet vegetable
Tajiki: Anganor with a crisp, crunchy texture and a sweet, nutty
Thailand: Thua-Duang flavour. The tubers are eaten raw or added to sal-
Turkish: Japon Enginari, Karabos, Cine Enginar ads or any cooked dish: stir-fries, casseroles,
Vietnamese: Trung Quốc Atisô soups, sweet and sour, sea food and curries.
Welsh: March Ysgall Tubers can be prepared in batter and fried as tem-
pura. They can be mixed with noodles, egg,
mushrooms or tofu. They can be steamed with
other vegetables like carrots, peas, zucchini and
Origin/Distribution onions. They can be steamed then tossed with
butter and herbs like chervil, parsley, savoury,
S. sieboldii is a native of China – Gansu, Hebei, chives and coriander or serve steamed artichokes
Hubei, Nei Monggol, Ningxia, Qinghai, Shaanxi, with a sauce, dressing or lemon juice. They can
Shandong, Shanxi, Sichuan and Xinjiang. It has be consumed with dips – avocado, kefir or cheese.
been cultivated as a food plant for its tubers since Tubers can also be pickled or dried, for later use.
ancient times in China and Japan. Chinese arti- In China, they are used mainly for pickling. In
choke referred to as Crosne in France has been Japan, the tubers are a part of Osechi-ryōri (tradi-
cultivated on a small scale since the end of the tional Japanese New Year Foods), which is
nineteenth century and in Malaysia (Cameron cooked to celebrate the Japanese New Year on 1st
Highlands) since the 1980s. It is occasionally January each year. In Japan, the tubers are pick-
planted in Mongolia, Korea, India, Great Britain, led in salt/vinegar, and purple perilla leaves are
Belgium, Switzerland, Germany, Italy, Russia, added for flavour and the red colour.
North America, Brazil, Argentina, South Africa
and New Zealand as a vegetable and medicinal
plant. Botany

A perennial, deciduous, erect or inclined herb,

Agroecology 30–120 cm high, hirsute throughout, stems quad-
rangular and soft, base furnished with short rhi-
Chinese artichoke is a cool climate temperate zomes thickened at the ends into fleshy, white,
species. In its native range, it occurs in lowlands, fusiform worm-like tubers, 3–5 cm long,
hillside and highlands from 0 to 3200 m altitude. 1–1.2 cm wide, slightly constricted at the nodes
It grows best in well-drained, moderately fertile, resembling a witchetty grub (Plates 1 and 2).
friable sandy soil in full sun. Excessive fertilisers Leaves opposite, green, vary in shape and size,
promote leaf growth and reduce tuber produc- ovate-cordate to ovate oblong, 2.5–9.5 cm by
tion. For optimum vegetative growth, a day tem- 1.5–3.5 cm wide, acute to shortly acuminate,
perature of 23 °C is required and for tuberization base cordate, serrate margin, petioles of leaves at
15 °C. It needs to be watered well during dry the middle of the stem 1–2 cm long, becoming
weather. The whole plant dies back in autumn, gradually shorter towards the stem apex. Flowers
which is when harvest occurs. rose purple, in terminal spicate panicles 10–15 cm
Stachys affinis 43

Plate 1 Fresh worm-like crosne


Plate 2 Dried crosne tubers

long, verticils of 6-flowers, subsessile; lowest

verticil subtended by lanceolate sessile leaves. Nutritive/Medicinal Properties
Calyx turbinate, 6–7 mm long, 5 toothed, teeth
subequal, ovate, acute, corolla 12–13 mm long, Tuber Phytochemicals
tube 9 mm long, limb bilabiate, upper lip arched,
hairy, lower lipped 3-lobed, reflexed, stamens 4, Tetraploid S. sieboldii was found to be more
ascending, anthers ovoid, nutlets 4 obovoid, dark superior in economic value than the diploid S.
brown, 1.5 mm across, tuberculate. sieboldii as the former contain higher levels of
44 Lamiaceae

nutritional components (Kuan 2012). The dry Leaf/Aerial Parts Phytochemicals

matter content, soluble protein content and solu-
ble sugar content increased 43.16 %, 9.46 % and From the aerial parts, the following glucosides
21.74 %, respectively, in the tetraploid leaf than were isolated: isoscutellarein 4’methyl ether
that in the diploid one; while they increased 7-O-B-(6”’-O-acetyl-2”-allosyl)glucoside, isos-
13.12 %, 21.89 % and 7.42 %, respectively, in the cutellarein 7-O-β-(6”’-O-acetyl-2”-allosyl)glu-
tetraploid tuber than that in the diploid one. coside and aceteoside (Takeda et al. 1985). Three
Tubers are rich in the oligosaccharide (tetra- new phenethyl alcohol glycosides together with
saccharide) carbohydrate – stachyose (α-galactose six known compounds stachysoside, acteoside,
(1→6) α- galactose (1→6) α-glucose (1→2) decaffeoylacteoside, isoacteoside, leucoscepto-
β-fructose), which is stored exclusively in the side A and martynoside were isolated from
vacuoles in the tubers (Keller and Matile 1985; Stachys sieboldii leaves (Nishimura et al. 1991).
Keller 1992b; Greutert and Keller 1993). The structures of three new compounds named
Vacuoles of Stachys sieboldii tubers were stachysosides A, B and C were established as
reported to accumulate up to 180 mM stachyose 2-(3,4-dihydroxyphenyl)ethyl O-α-l-arabino-
against a concentration gradient, probably by pyranosyl-(1→2)-α- l -rhamnopyranosyl-
means of an active stachyose/H+ antiporter situ- (1→3)-4- O -E-caffeoyl-β- d -glucopyranoside;
ated on the tonoplast (Greutert and Keller 1993). 2-(3,4-dihydroxyphenyl)ethyl O-α-l-arabino-
In dry tubers, 80 % of the dry weight comprised pyranosyl-(1→2)-α-L-rhamnopyranosyl-(1→3)-
stachyose. During an 18-day sprouting period, 4-O-E-feruloyl-β-d-glucopyranoside and 2-(3
the stachyose content decreased by 70 % and was -hydroxy-4-methoxyphenyl)ethyl O-α-l-arabino-
correlated with a twofold increase of the activity pyranosyl-(1→2)-α-l-rhamnopyranosyl- (1→3)-
of α-galactosidase, the key-enzyme responsible 4-O-E- feruloyl-β-d-glucopyranoside, respectively.
for the degradation of galactosyl saccharides.
Galactinol (gal-inositol), the precursor of galac-
tosyl saccharides, and sucrose were mainly pres- Antioxidant Activity
ent in the cytoplasm. Galactinol synthase (GS,
UDP-α-d-galactose: 1l-myo-inositol-1-O-α-d- Methanolic roots extracts of Stachys sieboldii
galactopyranosyltransferase) a key enzyme in the containing 3.02 % of polyphenols and 1.97 % of
biosynthetic pathway of the raffinose family of flavonoids exhibited strong antioxidant activities
oligosaccharides was found to be an extravacuo- (Baek et al. 2004). The fraction extracted by
lar enzyme in the tubers (Keller 1992a). The con- ethyl acetate showed higher antioxidation value
tent of stachyose in dry tubers was determined to than that of α-tocopherol, butylated hydroxyani-
be 236.0 mg/g, and the purity of the extracted sole (BHA) and butylated hydroxytoluene (BHT)
stachyose flour was calculated to be 87.34 % (Yin at the same concentration. The fraction ES-RS
et al. 2006). The tubers also contained sucrose exhibited the strongest activity on antioxidation
raffinose and verbascose (van den Burgh 1996; whereas the ES-R5 fraction showed similar pat-
Yin et al. 2006). Stachyose being a kind of sugar tern to flavones.
is used as a bulk sweetener and has been reported
to be better for health than normal white sugar
and has some interesting functional oligosaccha- Antimicrobial Activity
ride properties for the food industry, the cosmetic
industry and the pharmaceutical industry. The Stachys sieboldii root extract exhibited antimi-
tubers also contained stachysoside A, B and C crobial activity in-vitro against food-borne bacte-
acteoside with six known glycosides (Miyase ria – Bacillus cereus with MIc value of 250 μg/
et al. 1990); acteoside (1000 mg), stachysoside B ml, Listeria monocytogenes MIC =250–500 μg/
(280 mg), stachysoside C (50 mg), martynoside ml, Staphylococcus aureus and Psedomonas
(150 mg), leucosceptoside A (630 mg) and aeruginosa MICs of 500 μg/ml (Ryu and Park
phenylethanoid glycoside (Yamahara et al. 1990). 2002).
Stachys affinis 45

Anti-Anoxia Activity production against rabbit gamma-globulin in the

plasmas were lower than those of the nephritic
Studies using mice models also showed that control rats. Acteoside also suppressed hyper-
methanolic extract of the tubers had anti-anoxia cellularity and the incidence of crescent forma-
effect (Yamahara et al. 1990). The extract signifi- tion, adhesion of capillary wall to Bowman’s
cantly inhibited the lethal anoxia induced by capsule and fibrinoid necrosis in the glomeruli.
potassium cyanide (KCN) in mice. The methano- Furthermore, rat-IgG and C3 deposits on the
lic extract on fractionation yielded bioactive con- GBM were significantly less in the ACT-treated
stituents, acteoside and stachysoside C, group than in the control nephritic group.
phenylethanoid glycoside, which had a signifi- Acteoside also suppressed the up-regulation of
cant effect on the KCN-induced anoxia model. ICAM-1 expression in nephritic glomeruli and
prevented the up-regulation of ICAM-1 expres-
sion mediated by inflammatory cytokines or
Antitumour Activity phorbol 12-myristate 13-acetate on HUVECs
and rat mesangial cells. The results suggested
Administration of S. sieboldii ethanol extract to that acteoside may be a useful medicine against
mice accelerated mouse spleen cell growth but rapidly progressive glomerulonephritis, charac-
inhibited FM3A/S°-breast cancer cell growth terized by severe glomerular lesions with dif-
(Ryu et al. 2002). S. sieboldii fed mice showed a fuse crescents.
significant enhancement of interleukin IL-2
receptor expression, increased numbers of CD4+
T cells and CD8+ T cells. The extract also stimu- Central Nervous System Activity
lated the production of NO by peritoneal macro-
phages and spleen cells. The extract also inhibited Administration of Stachys sieboldii methanolic
Bl6Fl0 lung melanoma cells. extracts in the diet of rats for 20 days caused sig-
nificant inhibitory effects on acetylcholine
esterase activity, monoamine oxidase and xan-
Antinephritic Activity thine oxidase activities in the rat brain tissues
(Ryu and Kim 2004). Lipid peroxide levels were
Acteoside from S. sieboldii roots was found to also decreased in a dose-dependent manner.
have a suppressive effect on the accumulation of
leukocytes in the nephritic glomeruli through
prevention of the upregulation of adhesion mol- Traditional Medicinal Uses
ecules (Hayashi et al. 1994a, b, 1996). Acteoside
administered p.o. for 5 or 15 consecutive days The entire plant has been used in the treatment of
markedly suppressed the accumulation of total colds and pneumonia in traditional medicine in
leukocytes, ED-1-positive cells (monocytes/ China (Li and Hedge 1994). The dried and pow-
macrophages), CD4-positive cells, CD8- dered root is regarded as an anodyne (Duke and
positive cells, interleukin-2-receptor-positive Ayensu 1985).
cells (activated T cells) and Ia-positive cells in
the glomeruli of rats with crescentic-type anti-
glomerular basement membrane (GBM) nephri- Other Uses
tis. Acteoside inhibited the elevation of protein
excretion into urine. In acetoside-treated rats, The plant is used mainly as a vegetable crop with
cholesterol and creatinine contents and antibody incidental medicinal uses and no other uses.
46 Lamiaceae

Comments Keller F, Matile P (1985) The role of the vacuole in stor-

age and mobilization of stachyose in tubers of Stachys
sieboldii. J Plant Physiol 119(4):369–380
The plant is readily propagated from tubers. Kuan Q (2012) Quality analysis on tetraploid and diploid
of Stachys sieboldii Miq. Med Plant 3(9):71–73
Larkcom J (1991) Oriental vegetables. John Murray
Publishers Ltd., London. 232 pp
Selected References Li XW, Hedge IC (1994) Lamiaceae Lindley. In: Wu ZY,
Raven PH (eds) Flora of China, vol 17, Verbenaceae
Baek HS, Na YS, Kim DH, Lee CH, Ryu BH, Song SG through Solanaceae. Science Press/Missouri Botanical
(2004) Antioxidant activities of Stachys sieboldii Miq. Garden Press, Beijing/St. Louis. 378 pp
roots. Kor J Life Sci 14(1):1–7 Miyase T, Ueno A, Kitani T, Kobayashi H, Kawahara Y,
Duke JA, Ayensu ES (1985) Medicinal plants of China. Yamahara J (1990) Studies on Stachys sieboldii Miq.
Reference Publications, Inc., Algonac I. Isolation and structures of new glycosides. Yakugaku
Facciola S (1990) Cornucopia. A source book of edible Zasshi 110:652–657
plants. Kampong Publ., Vista, 677 pp Nishimura H, Sasaki H, Inagaki N, Chin M, Mitsuhashi H
Greutert H, Keller F (1993) Further evidence for stachy- (1991) Nine phenethyl alcohol glycosides from
ose and sucrose/h + antiporters on the tonoplast of Stachys sieboldii. Phytochemistry 30(3):965–969
Japanese artichoke (Stachys sieboldii) tubers. Plant Ryu BH, Kim SO (2004) Effects of methanol extract of
Physiol 101(4):1317–1322 Stachys sieboldii Miq on acetylcholine esterase and
Hayashi K, Nagamatsu T, Ito M, Hattori T, Suzuki Y monoamine oxidase in rat brain. Kor J Food Nutr
(1994a) Acteoside, a component of Stachys sieboldii 17(4):347–355
Miq, may be a promising antinephritic agent (2): effect Ryu BH, Park BG (2002) Antimicrobial activity of the
of acteoside on crescentic-type anti GBM nephritis in hexane extract of Stachys sieboldii Miq. Leaf. Kor
rats. Jpn J Pharmacol 65(2):143–151 J Life Sci 12(6):803–811
Hayashi K, Nagamatsu T, Ito M, Hattori T, Suzuki Y Ryu BH, Bg P, Song SK (2002) Antitumor effects of the
(1994b) Acteoside, a component of Stachys sieboldii hexane extract of Stachys Sieboldii Miq. Kor
Miq, may be a promising antinephritic agent (2): J Biotechnol Bioeng 17(6):520–524
effect of acteoside on leukocyte accumulation in the Takeda Y, Fujita T, Satoh T, Kakegawa H (1985) On the
glomeruli of nephritic rats. Jpn J Pharmacol glycosidic constituents of Stachys sieboldi Miq. and
66(1):47–52 their effects on hyaluronidase activity. Yakugaku
Hayashi K, Nagamatsu T, Ito M, Yagita H, Suzuki Y Zasshi 105(10):955–959
(1996) Acteoside, a component of Stachys sieboldii The Plant List (2013) Version 1.1. Published on the
Miq, may be a promising antinephritic agent (3): effect Internet;
of aceteoside on expression of intercellular adhesion van den Bergh MH (1996) Stachys sieboldii Miquel. In:
molecule-1 in experimental nephritic glomeruli in rats Flach M, Rumawas F (eds) Plant resources of South-
and cultured endothelial cells. Jpn J Pharmacol East Asia, No. 9, Plants yielding non-seed carbohy-
70(2):157–168 drates. Prosea Foundation, Bogor, pp 155–156
Hu SY (2005) Food plants of China. The Chinese Yamahara J, Kitani T, Kobayashi H, Kawahara Y (1990)
University Press, Hong Kong. 844 pp Studies on Stachys sieboldii MIQ. Anti-anoxia action
Keller F (1992a) Galactinol synthase is an extravacuolar and the active constituents. Yakugaku Zasshi
enzyme in tubers of Japanese artichoke (Stachys 110:932–935
sieboldii). Plant Physiol 99(3):125–1253 Yin J, Yang G, Wang S, Chen Y (2006) Purification and
Keller F (1992b) Transport of stachyose and sucrose by determination of stachyose in Chinese artichoke
vacuoles of Japanese artichoke (Stachys sieboldii) (Stachys sieboldii Miq.) by high-performance liquid
tubers. Plant Physiol 98(2):442–445 chromatography with evaporative light scattering
detection. Talanta 70(1):208–212
Maranta arundinacea

Scientific Name Vernacular Names

Maranta arundinacea L. Argentina: Jamaichipeke

Bolivia: Guate, Jamachipeke, Jamaichepeque,
Synonyms Brazil: Agoutiguepe, Agutiguepa, Araruta, Araruta-
comum, Araruta- palmeira (Portuguese)
Maranta arundinacea var. arundinacea, Maranta Cambodia: Daem Run, Saku
arundinacea var. indica (Tussac) Petersen, Chinese: Zhu Yu
Maranta arundinacea f. sylvestris Matuda, Costa Rica: Rizoma De Maranta, Sagú
Maranta arundinacea var. variegata Ridl., Cuba: Juá-Juá, Sagú De San Vicente
Maranta indica Tussac, Maranta ramosissima Wall., Czech: Maranta Rákosovitá
Maranta silvatica Roscoe, Maranta sylvatica Danish: Salepmaranta, Salepplante
Roscoe ex Sm., Maranta tessellata var. kegeljanii Dutch: Pijlwortel, Salepwortel
E.Morren, Phrynium variegatum N.E.Br. (illeg.) Eastonian: Roogmaranta
French: Arrowroot Des Antilles, Dictame
(Antilles), Dictame Barbed, Fécule De
Family Dictame, Herbe Aux Fleches
German: Maranta, Pfeilwurz
Marantaceae Guatemala: Shimipampana Blanco, Shimi-
pampana Negro, Silú, Sucu, Sulú, Tacea,
Common/English Names Hungarian: Bermudai Maranta, Maranta
India: Tikhor, Tikkor (Hindi); kuvehittu, tavak-
Arrow Head, Arrow-root, Arrowroot, Bermuda siri (Kannada), Kuva, kuvva (Malayalam),
Arrowroot, Maranta, Obedience Plant, St. Vincent Tavaksiri, Tugaksiri (Sanskrit),
Arrowroot, West Indian Arrowroot Aruruttukkilangu, Aruruttukkilangu, Kookai

© Springer International Publishing Switzerland 2016 47

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_4
48 Marantaceae

Neer, Koovai Kizhangu, Kuamau, Kukai Niru, Agroecology

Kuvai (Tamil), Palagunda, Palagunta (Telugu)
Indonesia: Angrik, Arus, Gaerut, Garut, Iruk, Arrowroot flourishes in warm, humid areas with
Jelarut, Larut, Nggarut, Rarut, Waerut mean annual temperatures of 25–28 °C and mean
(Javanese), Larut, Patat Sagu (Sundanese), annual precipitation of 1500–1800 mm. It thrives
Medawit Cina, Sugu Rarut (Sumatra) in well-drained, alluvial and volcanic soils and is
Japanese: Kuzuukon, Maranta also cultivated on yellow and brown latosols with
Laos: Sa:kh’u loamy-clayey texture as found in St. Vincent. It
Malaysia: Ararut, Berolu, Beruwi, Sagu Belanda, prefers partial shade as found in rainforest
Ubi bemban, Ubi Garut, Ubi Karut habitats.
Papiamento: Ararut
Peru: Saguero, Shimi pampana
Philippines: Sagu (Bikol), Galamaka (Bontok), Edible Plant Parts and Uses
Araru (Ibanag), Sagu (Iloko), Araru (Itogon),
Bai (Ivatan), Araro, Araru, Aroru, Aruru, Arrowroot rhizome and starch are used as food in
Uraro (Tagalog) the form of puddings, jellies, cakes, pastries, bis-
Portuguese: Agutiguepa, Araruta-comum, cuits, cookies hot sauces and used with beef tea,
Araruta-Especial, Araruta-palmeira, Aru-Aru milk, meal, broth and noodles in Vietnamese and
Puerto Rico: Juá-Juá, Maranta Korean cuisines. It provides easily digestible,
Russian: Maranta Trostnikovaia, Maranty nourishing food for children and people with
Trostnikovoi dietary restrictions and convalescents. It is well
Spanish: Ararú, Araruta, Caña Flecha, Chuchute suited as weaning food for infants from breast
Tamalera, Guape, Guate, Jamachipeke, milk. Arrowroot makes clear, shimmering fruit
Jamaichepeque, Juájuá, Juá-juá, Maranta, gels and prevents ice crystals from forming in
Rizoma De Maranta, Sagú, Sagú De San homemade ice cream. It can also be used as a
Vicente, Saguero, Silú, Sucu, Sulú, thickener for acidic foods, such as Asian sweet
Shimipampana, Tacea, Tamalera, Tubérculo and sour sauce. The absence of gluten in arrow-
De maranta, Yerén, Yuquilla root flour makes it useful as a replacement for
Swedish: Arrowrot wheat flour in baking.
Thailand: Sakhu (Central)
Turkish: Ararot
Uruguay: Tubérculo De Maranta Botany
Venezuela: Ararú, Araruta, Arrarú, Arrorúz,
Arrorruz, Arrurruz, Arruruz, Caña Flecha, A clumping perennial herb which is 30–130 cm
Chuchute Tamalera, Cuba, Guape, Guapo high with erect slender stems often apically
Vietnamese: Dong Củ branched (Plates 1 and 2). Rhizomes are white,
sympodial, cylindric, fleshy, starchy and loosely
scaly (Plates 5 and 6). Basal leaves are 4–8; cau-
line leaves 1–8; alternate, petiole 3.5–20 cm in
Origin/Distribution basal leaves, often absent in cauline leaves; pul-
vinus 0.2–1.8 cm, glabrous, adaxially slightly
Maranta arundinacea is native to South America, tomentose; lamina ovate-oblong, 3.5–35 by
the Caribbean and Mexico. The plant has natural- 3–11 cm, pale-green, adaxially sparsely pilose,
ized elsewhere in Florida, Australia, Southeast abaxially glabrous or sparsely pilose, base
Asia and South and East Africa. It is chiefly cul- rounded to truncate, apex acuminate (Plates 3
tivated in the West Indies (especially Jamaica and and 4). Inflorescences terminal, lax, several per
St. Vincent). leafy shoot, with 1–3 bracts each subtending 2–3
Maranta arundinacea 49

Plate 1 Young arrowroot plants Plate 2 Mature arrowroot clump

flower pairs, 2.5–6 cm. Flowers on common

pedicel; sepals green, narrowly ovate, corolla
white, tubular, base inflated, lobes 8–10 mm; sta-
minodes white, obovate, ovary three-loculed,
densely pubescent, to sub-glabrous. Fruit, green
or reddish-brown tinged, dehiscent, subglobose
capsule, 7–8 mm by 4–5 mm. Seeds brown,
rugose with basal aril.

Nutritive/Medicinal Properties
Plate 3 Large, ovate-oblong green leaves

Nutrients and Phytochemicals from Maranta arundinacea tuber had an amylose

content of 24.8 % (Madineni et al. 2012). The
Nutrient composition of the raw tuber per 100 g starch granules were small range of 2.92–
edible portion was reported as energy 125 cal, 6.42 μm, (mean of 4.84 μm), indented and spheri-
moisture 67.4 g, protein 1.7 g, fat 0.2 g, total car- cal, with length/degree of 1.20, and roundness of
bohydrates 29.5 g, dietary fibre 2.0 g, ash 1.2 g, 0.73. Maranta starch had a gelatinization tem-
Ca 15 mg, P 18 mg, Fe 1.9 mg, thiamin 0.13 mg, perature of 74.8°C, peak viscosity of 498BU and
riboflavin 0.02 mg, niacin 0.5 mg and ascorbic cold paste viscosity of 669 BU. It also possessed
acid 7 mg (Leung et al. 1972). Starch obtained higher freeze-thaw stability (15 cycles) 1.6,
50 Marantaceae

amylose 24.8 % and minerals (mg/100 g) such as

P 73.5 mg, Ca 17.2 mg, Mg 17 mg, K 27.4 mg, Fe
12.2 mg, Mn 10.9 mg and Zn 3 mg (Madineni
et al. 2012). In an earlier analysis, Perez and
Lares (2005) reported that arrowroot starch had
15.34 % moisture, 0.50 % crude protein, 0.18 %
crude fat, 0.47 % crude fibre, 0.21 % ash, 15.2 %
amylase, 84.79 % amylopectin and minerals
(mg/100 g) Na 57.26 mg, K 28.60 mg, P
75.10 mg, Ca 2.79 mg, Mg 12.24 mg, Fe 14.27 mg
and Zn 3.82 mg. Ensiled aerial biomass and
coarse and fine arrowroot processing residues
contained 10.8–21.1 % crude protein; 11.1–
30.2 % crude fibre; 3.8–17.0 % ash; and an in-
vitro dry matter digestibility of 38.5–60.3 %
(Erdman and Erdman 1984).
Jyothi et al. (2009) found that extrusion cook-
ing of arrowroot starch resulted in products with
very good expansion, colour and lower digestibil-
ity, which can be exploited for its potential use as
a snack food. The expansion ratio of arrowroot
starch extrudates ranged from 3.22 to 6.09 The
water absorption index (6.52–8.85 g gel/g dry
sample), water solubility index (15.92–41.31 %)
Plate 4 Arrowroot with variegated leaves and oil absorption index (0.50–1.70 g/g) were
higher for the extrudates in comparison to native
starch (1.81 g gel/g dry sample, 1.16 % and
0.60 g/g, respectively). The rheological proper-
ties, storage modulus and loss modulus of the
gelatinized powdered extrudates were signifi-
cantly lower and these behaved like solutions
rather than a paste or a gel. Hardness and tough-
ness were more for the samples extruded at
higher feed moisture and lower extrusion
temperature, whereas snap force and energy were
higher at lower feed moisture and temperature.
There was a significant decrease in the percent-
age digestibility of arrowroot starch (30.07 %
Plate 5 Immature young arrowroot rhizome after 30 min of incubation with the enzyme) after
extrusion (25.27–30.56 %).
Modification of native arrowroot starch with
swelling power at 30 °C =2.4 g/g, swelling power butyric anhydride increased swelling power and
at 80 °C 12.1 g/g, solubility at 30 °C was 6.3 % paste clarity, but decreased viscosity and solubil-
and solubility at 80 °C was 37.8 %. Arrowroot ity (Haryadi et al. 2008). Butyrylated arrowroot
starch contained moisture 8.1 %, crude protein starch at degree of substitution (DS) 0.04 at 2 %
0.8 %, crude fat 1 %, ash 1.5 %, soluble fibre (w/v) gave edible film of 0.05 mm thickness,
1.7 %, insoluble fibre 2.9 %, starch 81.6 %, sugar 0.32 MPa tensile strength, 1.66% elongation,
after inversion 1 %, sugar before inversion 0.2 %, 16.4 g/m2/h water vapour transmission rate
Maranta arundinacea 51

Plate 6 Mature arrowroot


(WVTR) and highest film solubility (46.8 %), cyclopenta-2,4-dienone; 2,3-dimethoxy-succunic

which completely dissolved after 1.40 min dip- acid dimethyl ester; 5-diethylsilanyloxy-4-ethyl-
ping in boiling water. The films could be applied 2-phenyl-3a,4,7,7a-tetrahydro-isoindole-1,3-dio
to provide instant arrowroot food powder. ne; triethyl-3-(3-methyl sulfanyl-1-vinyl-pent-1-
Dual modification of arrowroot starch using enyloxy)-silane; (2-methyl-thiiranyl)-methanol;
hydroxypropylation and cross-linking was found 2-tert-butoxy-tetrahydro-furan; cis-2(7-octynyl)
to overcome the lack of native arrowroot starch in cyclohexanol; 4-tert-butyl-[1,3,2]dioxathiolane-
food processing application (Maulani et al. 2-oxide; (2-acetoxy-1-methyl-vinyl)-methylidyne-
2013). These modifications significantly affected ammonium; tetradecane; cyclohepta-2,4,6-trie-
the composition of the amylose and amylopectin necarboxyclic acid ethyl ester; benzyl-butyl-
and the amount of phosphorus in the granules. amine; 1-ethoxymethyl-4-methyl benzene;
Higher amounts of phosphate salt gave a higher 1-(4-methoxy-cyclohexyl)-hex-5- en-1-one;
phosphorus content, which increased the degree 9-(4-methoxy-phenyl)-9-oxo-nonanoic acid methyl
of substitution (DS) and the degree of cross link. ester; 2,6-dimethoxy phenol; 2-tert-butyl-1,2-
Arrowroot starch that was modified using a con- dimethyl-cyclopropane, carboxylic acid methyl
centration of 8–10 % propylene oxide and 1–2 % ester; 1,1,2,2-tetramethyl-3-oxo-octahydro-4-
STMP (sodium trimeta phosphate): 3–5 % STPP oxa-cyclobuta(α)naphthalene- 2a- carbonitrile;
(sodium tri poly phosphate) produced a starch C-[2,2-dimethyl-3-(2-methyl-propenyl)-1-
with <0.4 % phosphorus content. A higher con- phenylsulfanyl- cyclopropyl]-methylamine;
centration of propylene oxide provided a higher 2-phenoxysulfonyl-acetimidic acid methyl ester;
degree of hydroxypropyl. 2-phenoxysulfonyl-acetimidic acid methyl ester
The ethanolic extract of M. arundinacea tuber hydrochloride; 3,6,10-trimethyl-8,11-dihydro-
exhibited a total phenol content of 390 mg GAE 7H-cyclodeca[b]furan-4-one;1-benzyl-4-tert-butyl-
(gallic acid equivalent)/100 g, total flavonoid 4,5-dihydro-1H[1,2,3,4,5]-thiatetrazoborole;
content of 290 QE (quercetin equivalent)/100 g 4-ethoxymethylene-7,7-dimethyl-bicyclo[3.2.0]
and total flavonol content of 150 mg QE/100 g. hept-2-en-6-one; 2-(2-nitroallyl)-cyclohexanone;
(Nishaa et al. 2013a). Also present were uniden- 1,4,7,10,10-pentamethyl-2,4,6,8,9-pentaaza-
tified steroid, tannin and glycoside compounds. t r i cy c l o [ 5 . 2 . 1 . 0 2 , 6 ] d e c - 8 - e n e - 3 , 5 - d i o n e ;
Forty-nine compounds were identified in the 2,3,3,4,7-pentamethyl-1,5,7-triaza- ricyclo
ethanol extract of arrowroot rhizome (Nishaa [,4]octane-6,8-dione; 6-chloro-3,4,4a,5,6,
et al. 2013b): cyclohexanone; 2-hydroxy- 8a-hexahydro-2H-chromene;5-(1-chloro-1-methyl-
52 Marantaceae

ethyl)-3,5-dimethyl-cyclopent-2-enone; 7a-(2- (Kumalasari et al. 2012). In addition, the arrow-

methoxy-ethyl)-1-methyl-1,2,3,6,7,7a- root tuber extracts strongly enhanced interferon γ
hexahydro-inden-5-one; 3-3-(methoxy-phenyl)- production by splenocytes. In in-vivo study, the
2-methyl-oxetan-3-ol; 2-benzyloxy-7-(tetrahydro- diet containing arrowroot extracts increased the
pyran-2-yloxy)-heptan-1-ol; (1-acetyl-5-formyl- serum IgG, IgA and IgM levels in mice.
6-methyl-cyclohexa-2,4-dienyl)-acetic acid ethyl
ester; 3-phenyl-1-(toluene-4-sulfonyl)-pyrrolidine-
2,5-dicarboxylic acid 2-benzyl ester 5-tert-butyl Anti-Diarrhoeal Activity
ester; cyclopropyl-oxo-acetic acid methyl ester;
2-allyl-5a-hydroxy-octahydro-5-oxa- In a small clinical study of 11 patients with diar-
2-aza-cyclopenta[c]inden-1-one;cyclohexylmethyl- rhoea, the administration of arrowroot powder
diethyl-methoxy-silane; 2,2-dimethoxy-4a,5,6, (thrice daily) for a month was found to be an
7,8,8a-hexahydro- 2H-benzo[e][1,2]oxasilane; effective treatment for diarrhoea (Cooke et al.
2,2-dimethoxy- 2H-benzo[e][1,2]oxasilane; 2000). Arrowroot reduced diarrhoea and had a
4-(3,4-dimethoxy-phenyl)-butan-1-ol; long-term effect on constipation. It also eased
1,1,diethoxy-2-methyl-propane; 2,4’-dimethyl- abdominal pain. Its action was attributed to an
[2,4’]bi[[1,3]dioxanyl]; 2-methyl-3,3-bis-(2- increase in faecal bulk leading to more efficient
trimethylsilane-ethoxy)-propionic acid methyl bowel action.
ester; 2-methoxyimino-4-methyl-pentanoic acid
benzyl ester; 2-(benzyl-{2-[(dimethyl carbam-
oyl-phenyl-methylene)-hydrazino]-ethyl}- Probiotics Enhancing Activity
and 3-methylene-1-oxa-spiro[3,6]decane. Studies showed that arrowroot carbohydrates
could be used to enhance probiotic Lactobacilli
population in bio-yoghurt during refrigerated
Antioxidant Activity storage (Abesinghe et al. 2012). Arrowroot car-
bohydrates and Raftilose® increased Lactobacilli
The ethanolic extract of M. arundinacea rhizome population by 1.44 log CFU/mL and 1.17log
exhibited high antiradical activity against CFU/mL, respectively, compared to the
1,1-diphenyl-2-picryl hydroxyl (DPPH) ABTS control.
acid), hydrogen peroxide and nitric oxide radicals
with IC50 value of 293.4, 297.4, 336.1 and Antibacterial Activity
258.7 μg/ml, respectively (Nishaa et al. 2012).
The reducing power and ferric reducing antioxi- Studies by Kim and Fung (2003) found that
dant power (FRAP) increased with increasing 0.63 % of arrowroot tea was effective in inhibit-
concentration of the sample. The antioxidant ing four food-borne pathogenic bacteria,
activity of the sample was comparable with that of Escherichia coli, Salmonella enteritidis, Listeria
the standard butylated hydroxyl toluene (BHT). monocytogenes and Staphylococcus aureus to the
minimum detection limit of 1 log cfu/ml at 5 day
in brain heart infusion (BHI) medium.
Immunostimulatory Activity

Studies showed the arrowroot tuber extract stim- Enterokinase Acitivty

ulated IgM production by human hybridoma
HB4C5 cells and immunoglobulin (IgG, IgA and Maranta arundinacea tuber exhibited endoge-
IgM) production by murine splenocytes in-vitro nous esterase activity towards benzoyl arginine
Maranta arundinacea 53

ethyl ester (Bhat et al. 1981). Enterokinase is a Comments

serine proteinase required for the activation of
pancreatic proenzymes in the duodenum. Arrowroot can be propagated by suckers and
rootstock or rhizomes.

Traditional Medicinal Uses

Selected References
Traditionally the tuberous rhizomes are used in
the treatment of diarrhoea (Nishaa et al. 2012). Abesinghe N, Vidanarachchi J, Silva S (2012) The effect
of arrowroot (Maranta arundinacea) extract on the
Arrowroot is mainly useful as an easily digested,
survival of probiotic bacteria in set yoghurt. Int J Sci
nourishing diet for convalescents especially for Res Pub 2(5):1–4
bowel complaints as it has demulcent properties Anonymous (2002) Ethanol from arrow root (Maranta
(Grieve 1971). Arrowroot starch in jelly form is arundinacea Linn) SCINET Phil (S-14). 29 p. http://
suitable as weaning food for infants. Mashed rhi-
zomes are applied topically for wounds from poi- w+root+%28Maranta+Arundinacea+Linn%29&Mtyp
soned arrows, scorpion and black spider bites and e=PROJECTS
to arrest gangrene. Freshly expressed juice mixed Bhat PG, Jacob RT, Pattabiraman TN (1981) Enzyme
inhibitors from plants: Enterokinase inhibitors in
with water is used as antidote internally for veg-
tubers and seeds. J Biosci 3(4):371–378
etable poisons. Cooke C, Carr I, AbramS K, Mayberry J (2000) Arrowroot
as a treatment for diarrhoea in irritable bowel syn-
drome patients: a pilot study. Arq Gastroenterol
Other Uses Erdman MD, Erdman BA (1984) Arrowroot (Maranta
arundinacea), food, feed, fuel, and fiber resource.
Arrowroot was found to be a food, feed, fuel Econ Bot 38(3):332–341
and fibre resource (Erdman and Erdman 1984). Grieve M (1971) A modern herbal. Penguin. 2 vols. Dover
publications, New York, 919 pp
Theoretical yields of 0.27 and 1.60 l of methane
Haryadi, Kusumasmarawati AD, Marseno DW (2008)
at standard temperature and pressure per litre of Application of butyrylated arrowroot-starch for edible
arrowroot rhizome wash water and starch-set- film to contain instant food powder. In: Proceedings
tling water were calculated, respectively. Fuel the 1st conference on food science and technology,
Mekong Delta, Can Tho University, Vietnam. March
alcohol production potential from yeast-supple-
20–22, 2008, pp 47–50
mented aerial biomass and coarse residue of Jyothi AN, Sheriff JT, Sajeev MS (2009) Physical and
arrowroot were identified. Arrowroot coarse functional properties of arrowroot starch extrudates.
residue had qualities that may be suited to tear- J Food Sci 74:E97–E104
Kim S, Fung DYC (2003) Antimicrobial effect of water
resistant specialty grade papers, such as wrap-
extract of arrowroot (Maranta arundinacea) tea on
ping paper and bags. Studies involving foodborne pathogens in liquid medium. Session 29F,
production of extract, fermentation and distilla- Food microbiology: control of foodborne microorgan-
tion processes conducted in the Philippines isms by antimicrobials IFT (Institute of Food
Technologists) annual meeting, 12–16 July 2003,
reported arrowroot rhizome to be a promising
source of ethanol with 56 % purity (Anonymous Kumalasari ID, Harmayani E, Lestari LA, Raharjo S,
2002). In remote Philippine barrios, arrowroot Asmara W, Nishi K, Sugahara T (2012) Evaluation of
starch is also used for starching clothes (Stuart immunostimulatory effect of the arrowroot (Maranta
arundinacea. L) in vitro and in vivo. Cytotechnology
Arrowroot can be used as screens between Leung WTW, Butrum RR, Huang Chang F, Narayana Rao
neighbouring houses. It provides excellent com- M, Polacchi W (1972) Food composition table for use
post and mulch. Shoot tips and leaves can be uti- in East Asia. FAO, Rome. 347 pp
Madineni MN, Faiza S, Surekha RS, Ravi R, Guha M
lised as feed for pigs, goats and chickens. The
(2012) Morphological, structural, and functional prop-
individual leaves can be used as plates during erties of maranta (Maranta arundinacea L) starch.
picnics. Food Sci Biotechnol 21(3):747–752
54 Marantaceae

Maulani RR, Fardiaz D, Kusnandar F, Sunarti TC (2013) Porcher MH et al. (1995–2020) Searchable world wide
Characterization of chemical and physical properties of web multilingual multiscript plant name database.
hydroxypropylated and cross-linked (Marantha arun- Published by The University of Melbourne, Australia.
dinacea) starch. J Eng Technol Sci 45(3):207–221
Nishaa S, Vishnupriya M, Sasikumar JM, Hephzibah PC, Frontpage.html
Goaplakrishnan VK (2012) Antioxidant activity of Stuart GU (2013) Philippine alternative medicine. Manual
ethanolic extract of Maranta arundinacea L. tuberous of some Philippine medicinal plants. http://www.stu-
rhizomes. Asian J Pharm Clin Res 5(4):85–88
Nishaa S, Vishnupriya M, Gopalkrishnan VK (2013a) The Plant List (2013) Version 1.1. Published on the
Qualitative assessment of ethnolic extract of Maranta Internet;
arundinacea L. rhizome using HPTLC. Int Res USDA, ARS (2013) National Genetic Resources
J Pharm 4(2):76–83 Program Germplasm Resources Information
Nishaa S, Vishnupriya M, Sasikumar JM, Gopalkrishnan Network – (GRIN) [Online Database], National
VK (2013b) Phytochemical screening and GC-MS anal- Germplasm Resources Laboratory, Beltsville,
ysis of ethanolic extract of rhizomes of Maranta arundi- Maryland.
nacea L. Res J Pharm Biol Chem Sci 4(2):52–59 html/
Pérez E, Lares M (2005) Chemical composition, mineral Wu D, Kennedy H (2000) Marantaceae. In: Wu ZY, Raven
profile, and functional properties of canna (Canna PH (eds) Flora of China, vol 24, Flagellariaceae
edulis) and arrowroot (Maranta spp.) starches. Plant through Marantaceae. Science Press/Missouri
Food Hum Nutr 60(3):113–116 Botanical Garden Press, Beijing/St. Louis
Nelumbo nucifera

Scientific Name Common/English Names

Nelumbo nucifera Gaertner Baladi Bean, Bean Of India, Chinese Water Lily,
East Indian Lotus, Egyptian Bean, Hindu Lotus,
Indian Lotus, Lotus, Lotus Bean, Oriental Lotus,
Synonyms Sacred Lotus, Sacred Water Lotus, Water Lotus

Nelumbium album Bercht. & J.Presl, Nelumbium

asiaticum Rich., Nelumbium caspicum Fisch. ex Vernacular Names
DC., Nelumbium caspium Eichw., Nelumbium
discolor Steud., Nelumbium indicum Poir.,
Nelumbium javanicum Poir., Nelumbium margin- Arabic: Karambeulma, Kanwal Gatta, Nilufer,
atum Steud., Nelumbium nelumbo Druce, Ussulnilufer
Nelumbium nuciferum Gaertn., Nelumbium rhee- Brazil: Lotus
dii C.Presl, Nelumbium speciosum Willd., Burmese: Padung Ma
Nelumbium tamara Sweet, Nelumbium transver- Chinese: Lian, Lien (Plant), Ou (rhizome), He-
sum C.Presl, Nelumbium turbinatum Blanco, Hua, Ho-Hua (flower), Lian Zi, Lien Tzu
Nelumbium venosum C.Presl, Nelumbo caspica (Seed), He-Ye, Ho-Yeh (Leaf), Lien-Yong,
(Fisch.) Schipcz., Nelumbo caspica Fisch., Lien Yung (Mashed Cooked Lotus Seed), Ou-
Nelumbo indica Pers., Nelumbo komarovii Fen (Dried Rhizome Starch)
Grossh., Nelumbo nelumbo (L.) H.Karst., Czech: Lotos Indický
Nelumbo nucifera var. macrorhizomata Nakai, Danish: Indisk Lotus, Lotus
Nelumbo speciosa Willd., Nelumbo speciosa var. Dutch: Indische Lotusbloem, Lotus
alba F.M.Bailey, Nymphaea nelumbo L., Tamara Estonian: India Lootos
alba Roxb. ex Steud., Tamara hemisphaerica French: Fève d’Egypte, Lotier Lotus Sacré,
Buch.-Ham. ex Pritz., Tamara rubra Roxb. ex Lotus Égyiptien, Lotus Indien, Lis Du Nil,
Steud. Lotus Des Indes, Lotus Magnolia, Racines De
Lotus, Rose Du Nil
German: Agyptische Bohne, Indische
Family Lotosblume, Indischer Lotos, Indischer Lotus,
Lotos, Lotosblume, Lotus, Nilli Lili, Padma
Nelumbonaceae Hungarian: Indiai Lótusz (Virág)

© Springer International Publishing Switzerland 2016 55

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_5
56 Nelumbonaceae

Indonesia: Patma, Tarate (Malay), Tarate, Trate Cattalam, Catti, Cattika, Cattikakkoti,
(Javanese), Tarate (Sundanese) Caturaccakan, Cekkamalar, Cenkalam,
India: Podum (Assamese), Kamal, Kombol, Centamarai,Ceppilai, Ceyyatamarai, Cirukam,
Komal, Padama, Padma, Pudmapudu Cirukam, Citalam, Citampocam, Citampucam,
(Bengali), Ambuj, Kamal, Kamal-Kakri, Cittetan, Civacattipakirtan, Comakkiyam,
Kanwal, Kanval, Lalkamal, Padam (Hindu), Copanam, Cuceyam, Cultalai, Cultalai,
Kamala, Tavare, Tavaribija, Tavarigadde, Curiyanatpu,, Elimanai, Ellimanai, Intai,
Thaavare (Kannada), Bem-Tamara, Iracacupakkoti, Iracacuyam, Iracalam,
Bemtamara, Centamara, Tamara, Ventamara Iraciyam, Iramapiriyam, Iramappiriyam,
(Malayalam), Thambal (Manipuri), Kamal, Iramappiriyam, Iramappiriyam, Irampu,
Kamala (Marathi), Kamal (Oriya), Abja, Iratacanniyakam, Iratikantal, Iratikantan,
Ambhoja, Ambhoruha, Amboruha, Ambuja, Irattacantiyakam, Irattacarorukam,
Ambujanma, Ambupadma, Amburoha, Irattakamalam, Irattakkumutam,
Amlana, Aranala, Aravinda, Arvinda, Irattakokanam, Irattakumutam, Irattalamutam,
Bisakusuma, Bisaprasuna, Drishopadma, Irattamantalam, Irattorpalam, Iravikantam,
Harivetra, Indiralaya, Jalajanma, Kalhara, Irciyam, Jalacam, Kacam, Kamalam,
Kamala, Kamalam, Kamalodbhavamrajah, Kamalini, Kancam, Kancamalar, Kancankoti,
Kanval, Kawar, Kunja, Kusesaya, Kusheshaya, Kantokakkoti, Kantokam, Kantotam, Karkam,
Kutapa, Mahapadma, Mahotpala, Nala, Katirppari, Kauravakkoti, Kauravam,
Nalika, Nalina, Padma, Padmadrajah, Kokanakam, Kokanakam, Kokayam,
Padmam, Pankaj, Pankaja, Pankajam, Makorpalakkoti, Makorpalam, Malunti,
Pankeruha, Pathoja, Pundarika, Pundra, Maraikkoti, Matanalam, Mirinalam,
Pushkara, Puskara, Putaka, Rajiva, Mirunalakkoti, Mirunalam, Mirunali,
Sahasrapatra, Sahsrapatra, Sarasa, Sarasiruha, Mirunarakkoti, Mirunaram, Mirunarram,
Sarojanma, Saroruha, Sarsija, Sarsiruha, Mirutalakkoti, Mirutalam, Muntakakkoti,
Satapatra, Sharada, Sharapadma, Shatapatra, Muntakam, Nalaki, Nalaki, Nalakikkoti,
Shri, Shriparna, Shrivasa, Shuklapadma, Nalalmatu, Nalam, Nalikam, Nalini, Nalitam,
Sitambuja, Sujala, Svetakamala Tamarasa, Nallatamarai, Naticam, Naticam,
Tumarasa, Vanashobhana, Varisoha, Tra Nattuttamarai, Nicakaci,
(Sanskrit), Acaiyapattipam, Acaiyappattiram, NicakacikkotiNirmelcevvanti, Nirmelpaccilai,
Aciyapattiarm, Akiyapattiram, Allakam, Nirnatittamarai, Nirniti, Nirorukakkoti,
Ambal, Ampala, Ampo, Ampocam, Nirorukam, Palututainayakam, Panikam,
Ampocayoni, Ampucam, Ampucani, Panikkanci, Panikkancikkoti, Pankacam,
Ampucanmam, Ampucatam, Ampucini, Pankacatam, Pankacatamakkoti,
Ampunecam, Ampuracam, Ampurecam, Pankacatamam, Pankarukam, Pankatam,
Ampuru, Amputam, Ampuyakkoti, Ampuyam, Pankecam, Pankeram, Panmam, Pannaci,
Ampuyatam, Anikini, Anikini, Appucam, Parparakam, Patmam, Patocam, Patotam,
Arantam, Arappatumam, Arpakantam, Patumakam, Patumanalakkoti, Ponmanai,
Arpatumam, Arukanvakanam, Arunakamalam, Potu, Poykainari, Poykainarinir, Puntaram,
Arunkalaccevvital, Aunikam, Ayamalar, Puntarikam, Putpan, Rattorpalam, Tamaracam,
Aymalar, Caccatam, Calacanam, Calacanmam, Tamaram, Tantamarai, Tantar, Tantulam,
Calacappu, Calacarakkuli, Calakankakkoti, Tanturam, Tarutam, Tevanam, Tikkayam,
Calakankam, Calakarankam, Calarukakkoti, Tirumalarkkoti, Tirumalkompar, Tirumalunti,
Calarukam, Calcam, Calilikam, Cantiram, Totakam, Toyacam, Tunaparicam,
Caracakkoti, Caracam, Caracijam, Sivapputamarai, Tamarai, Thamarai,
Caracirukam, Caravanam, Carocini, Thaamarai, Tamaray, Ventamarai
Carokkam, Carokulam, Catalam, Catapattiram, Vicappicuranam, Vicappiracunam,
Catapatumam, Catippakitakkoti, Catippakitam, Vicaputpam, Vintu, Vintukam Visaputpam,
Nelumbo nucifera 57

Urocanai, Urocani, Urokani, Vacanakkoti, New Guinea, Pakistan, Philippines, Russia (Far
Vacanam, Vanacakkoti, Vanacopanam, East), Sri Lanka, Thailand, Vietnam, Laos,
Vanicam, Vannivannam, Vantuni, Vantunikkoti, Kampuchea and northern Australia.
Varicam, Varicam, Varikam, Vaucikakkoti,
Vaucikam (Tamil), Damara, Erratamara,
Errathaamara, Kalung, Kamalam, Kamalamu, Agroecology
Padmamu, Pundreekamu, Soungadhikamu,
Tamara, Thaamara, Thaamara Puvvu, Thella Lotus is strictly aquatic. It grows wild and is culti-
Thaamara, Thellane Padmanu (Telugu), vated in the mud of shallow, sheltered waters of
Nilufer (Urdu) lakes, ponds and inundated rice fields from sea level
Italian: Fior Di Loto, Giglio De Nilo, Loto to 1500 m elevation in the tropics and subtropics.
d’Egitto, Ninfea d’Egetto
Japanese: Hasu, Hasu-N-Né, Hasu-No-Mi,
Renkon Edible Plant Parts and Uses
Khmer: Chhuk
Korean: Yeongeun, Yon Puri (Rhizome) Lotus is well known as a food and medicinal
Laotian: Bwà plant, and its every part is utilised. All parts of the
Malaysia: Bunga Padam, Bunga Telepok, lotus plant, including flowers, seeds, leaves,
Padema, Seroja, Teratai stems and roots are consumed (Sridhar and Bhat
Nepal: Kamal, Raato Kamal, Raato Thuulo 2007) (Plate 7). In Korea, lotus leaf, root and
Kamal seed are usually consumed as a tea, or in braised
Persian: Beykhneelufer, Nilufer, Nilufu dishes or soups (Ha et al. 2010). Tender rhizomes,
Philippines: Liñgaling (Ibanag), Sukau (Iloko), stems and leaves of lotus are edible and can be
Saua (Maguindanao), Baino (Tagalog) cooked along with other vegetables, soaked in
Portuguese: Flor-De- Lótus, Lótus Do Egipto, syrup or pickled in vinegar (Phillips and Rix
Lótus-Do-Egito, Lótus Índico, Lótus-Sagrado, 1993). Rhizomes have mild flavour and are exten-
Sementes De Lótus sively used in Chinese recipe, while stem is used
Spanish: Habas De Egipto, Loto Sagrado, Rosa in cooking as food and it tastes like beet (Hedrick
Del Nilo, Semillas De Loto, Nelumbio Raiz 1972; Tanaka and Nguyen 2007). Ogle et al.
De Lotus (Root) (2001) reported the use of lotus stem (consisting
Sri Lanka: Nelum (Sinhalese) of 6, 2.4 and 0.2 mg/100 g calcium, iron and zinc,
Taiwan: Lian respectively) as a vegetable used in salads at
Thailand: Bua-Luang, Sattabut, Ubon Vietnam. The seeds can be popped like popcorn,
Tibetan: Pa Dma Dkar Po, Pa Dma Dmar Po, ground into powder and eaten dry or used in
U-Tpa-La bread making. The roasted seeds are good coffee
Vietnam: Hoa Sen, Sen, Lien, Ngau (Tay), Bo substitute and possess saponins. The products of
Bua (Thai), Lin Ngo (Dao) lotus rhizome, such as fresh, salted and boiled
lotus rhizome and lotus rhizome starch, drinks,
teas and lotus seeds, are very popular in the daily
diet (Hu and Skibsted 2007; Zhong et al. 2007).
Origin/Distribution Starches extracted from lotus rhizomes are com-
mercially available in China and consumed in
The species is distributed from the Caspian Sea fast food and as breakfast, traditional confection-
and Iran through Asia to northern Australian – ery and food additives (Zhong et al. 2007). The
China (predominantly in Zhejiang, Jiangsu, Korean traditional lotus liquor (Yunyupju) is
Anhui, Hunan, Hubei), Bhutan, India, Indonesia made from lotus blossom and leaves (Lee et al.
(Java), Japan, Korea, Malaysia, Myanmar, Nepal, 2005). Lotus leaves are used for tea in China
58 Nelumbonaceae

(Zhou et al. 2013a). Lotus leaves are used for tea seed soup dessert’ with longan aril seeds, which
or food like yeon yip bap in Korea (Ahn et al. can be served warm or cold with ice. The Chinese
2013). in China and elsewhere also use lotus seeds as a
The main types of lotus rhizome processed common ingredient in cooling ‘bupin’ mixtures.
products are the salt bloated lotus, deep frozen In India, the seeds after removal of the outer testa
lotus and vacuum-packed lotus (Guo 2009). All is consumed raw, or dried and puffed like
these products are mainly exported to Japan, fol- popcorn.
lowed by South Korean and other southeast Asian The roots (rhizomes) are a food used exten-
countries. Lotus is one of 26 vegetables with sively in China, Korea and Japan and in southeast
largest sale in China (Jiang and Cao 2005). Asia are sold whole or in cut pieces, fresh, frozen
Currently, the lotus herb is becoming more popu- or canned. It has a crunchy texture with sweet-
lar in China as a ‘tea drink’ or as a main ingredi- tangy flavours. The fresh rhizomes are commonly
ent of some herbal formulations (Ye et al. 2014a). sliced and eaten raw or cooked with meat in
Lotus leaf is used as food or beverage in Taiwan soups or fried with salad, prawns, meat, sesame
(Lee et al. 2010). In Sulawesi, the young shoots oil and/or coriander leaves in many Asian cui-
and unexpanded leaves are eaten boiled, and the sines. In Thailand, the young rhizomes are peeled
leaves may be eaten raw (Burkill 1966). The rhi- and eaten fresh in salads, while in China, Japan,
zomes are cooked as food or steam and eaten in Malaysia and elsewhere, the mature rhizomes are
salad. They are also pickled in salt or vinegar. stir-fried, stuffed and deep fried or cooked in
The unripe seeds are eaten raw, boiled or soups. Lotus rhizome is also common in Japanese
roasted, while the ripe seeds are boiled or roasted. ‘bento’ food boxes and ‘nimono’. In Korea, the
Lotus seeds are used in soups, food dishes, sweet- root is used as a vegetable in soups and deep-
meat, cakes, pastries, desserts and nourishing fried, stir-fried and braised dishes. Lotus roots
herbal ‘bupin’ tea. Dried seeds, sometimes called are often pickled with rice vinegar, sugar, chilli
‘lotus nuts,’ must be boiled until softens. The and or garlic. Dried rhizome pieces are also fried
seeds can be crystallised with sugar as part of as chips. A high-quality edible starch is obtained
Chinese New Year sweet offerings, cooked into a from the rhizomes and is used as baby food and
sweet soup called ‘tong sui’ together with dried as a special diet for healing sickness in China.
longan and rock sugar and mashed and cooked The young leaf petioles, after the rough, outer
into sweetened lotus nut paste (‘Lien-yong’) layer has been scraped off, can be found in local
which is used as a filling for Chinese moon cakes markets and are used in preparing salads and
(in mid-Autumn, it is customary to serve ‘moon soups or cooked as vegetable with other meat
cakes’ which have a filling made of lotus seeds dishes in Thailand, Vietnam and Indonesia
and walnuts), ‘daifuku’ (Japanese glutinous rice (Plate 9). The petioles are also slivered (Plate 10).
cake with sweet lotus past as fillings), rice flour In China and Indo-China, flower stamens are
pudding, pastries, buns, bread and sauces. The used to perfume herbal tea called ‘liánhuā cha’ in
seeds are used to prepare ‘tong sui’ (sweet soup). China and trà sen and chè sen in Vietnam.
The seeds can be cooked in soups, usually with Likewise in India, the stamens of the flowers
chicken or beans. An example of the latter is a infused with water are employed to make a fra-
‘red bean and lotus seed soup’ served at banquets grant tea. The dried lotus flowers are used in
for newlyweds, comprising red beans and lotus cooking dishes, such as ‘Mandarin duck and
seeds. Red beans (hong dou) represent strength, lotus flowers’, and the fresh petals are also edi-
while lotus seeds (lian zi) symbolise the newly- ble. In Thailand, young tender leaves are cooked
weds being blessed with a child each year. The and eaten with a savoury sauce, and the flower
soup is also presented at the Chinese New Year’s petals can be dipped in ‘nam prik’ or used as gar-
festival. Other popular and common dishes is the nish. Dried leaves are used for herbal tea in
‘cream lotus seed soup’ which is prepared with China. In Korea, the leaves and petals are used as
crushed pineapple pieces and the ‘sweet lotus a tisane: ‘yeonkkotcha’ made with dried petals of
Nelumbo nucifera 59

white lotus and ‘yeonipcha’ made with leaves. sparsely spinulate (Plates 1, 3). Tepals are cadu-
Lotus leaves are used as wrapper for Chinese cous, pink or white, oblong elliptic to obovate,
‘tamale’, a special Chinese pastry in China. The 5–10 × 3–5 cm and spirally arranged. Stamens are
leaves are used as flavouring and wrapping for numerous, slightly longer than receptacle; fila-
rice preparations in making dim sum and other ment is slender; anther is linear, 1–2 mm; and
sticky, glutinous rice preparations. Lotus leaves connective appendage is clavate, 7 mm long and
are also used for wrapping whole chicken, mari- incurved. Ovary is apocarpous, embedded in an
nated and douched with rice wine, soy sauce, obconical, spongy, accrescent, turbinate 5–10 cm
mushroom and sometimes with some other in diameter receptacle. Its fruit is a nut oblong to
Chinese herbs before the whole entity is encased ovoid, 1.0–2.0 × 7–1.5 cm and glabrous; pericarp
in mud or clay and cooked to give the much- is thick, hardened (bony) and brown (Plates 4, 5).
relished restaurant specialty called ‘beggars Seed is having a reddish brown testa, two yellow-
chicken’ or ‘drunken beggars chicken’. ish-white mealy cotyledons and a stalked green
embryo (Plates 5, 7, 8).

Nutritive/Medicinal Properties
Nelumbo nucifera is a perennial, rhizomatous,
aquatic herb with the horizontal creeping rhi- Seed/Pod/Receptacle Nutrient/
zomes and roots buried in the mud (Plate 1). The Phytochemicals
rhizome has nodes externally and forms roots at
these nodes, and internally it has aerenchyma, Analyses carried out in the United States reported
which is a parenchyma tissue with large intercel- that raw mature lotus seeds had the following
lular air spaces (Plate 6). The leaves are raised proximate composition (per 100 g value): water,
above the water or floating, on terete, fistulous, 77 g; energy, 89 kcal (372 kJ); protein, 4.13 g;
glabrous or prickly, hollow, 1–2 m long petiole total lipid, 0.53 g; ash, 1.07 g; carbohydrates,
(Plates 9, 10). Leaf’s blade is large, abaxially 17.28 g; Ca, 44 mg; Fe, 0.95 mg; Mg, 56 mg; P,
blue-green, orbicular, peltate, 25–90 cm in diam- 168 mg; K, 367 mg; Na, 1 mg; Zn, 0.28 mg; Cu,
eter, papery, glabrous, glaucous and water repel- 0.094 mg; Mn, 0.621 mg; vitamin C, 0 mg; thia-
lent, having margin entirely (Plates 2, 5). Flowers mine, 0.171 mg; riboflavin, 0.040 mg; niacin,
are large, showy, solitary 10–23 cm in diameter 10.429 mg; pantothenic acid, 0.228 mg; vitamin
on scape longer than petioles, glabrous or B-6, 0.168 mg; total folate, 28 μg; vitamin A,

Plate 1 Lotus flowers, fruit and

60 Nelumbonaceae

Plate 2 Large peltate, orbicular,

waxy leaf

Plate 3 Close up of fertilised lotus


13 IU; total saturated fatty acids, 0.088 g; 14:0

(myristic acid), 0.001 g; 16:0 (palmitic acid),
0.077 g; total monounsaturated fatty acids,
0.104 g; 18:1 undifferentiated (oleic acid),
0.062 g; 20:1 undifferentiated (eicosenoic acid),
0.012 g; 22:1 undifferentiated (erucic acid),
0.031 g; total polyunsaturated fatty acids 0.312 g;
18:2 undifferentiated (linoleic acid), 0.285 g;
18:3 undifferentiated (linolenic acid) 0.027 g;
tryptophan, 0.059 g; threonine, 0.200 g; isoleu-
cine, 0.205 g; leucine, 0.326 g; lysine, 0.264 g;
methionine, 0.072 g; cystine, 0.054 g; phenylala-
nine, 0.206 g; tyrosine, 0.100 g; valine, 0.266 g;
arginine, 0.338 g; histidine, 0.115 g; alanine,
0.239 g; aspartic acid, 0.505 g; glutamic acid, Plate 4 Lotus fruit
Nelumbo nucifera 61

Plate 5 Lotus fruit (TS) and peeled

an unpeeled fresh lotus seeds

Plate 6 (a, b) Harvested lotus roots

Plate 7 Lotus root (left) lotus seeds

and leaves sold in the market
62 Nelumbonaceae

Plate 8 Dried processed lotus seeds

Plate 9 Lotus leaf petioles, peeled

and used as food

0.957 g; glycine, 0.221 g; proline, 0.344 g; and

serine, 0.252 g (USDA-ARS 2014). The crude
protein of lotus seeds was found to be 14.81 %
(Ibrahim and El-Eraqy 1996). The essential
amino acids found were threonine, 2.43 %;
methionine, 0.82 %; leucine, 3.33 %; isoleucine,
1.11 %; and phenylalanine, 12.64 %. The semi-
essential amino acids were 7.45 % arginine and
10.79 % histidine. Other amino acids found in
adequate amount were glutamic, 26.56 %; pro-
line, 10.76 %; aspartic, 6.17 %; tyrosine, 4.65 %;
serine, 4.27 %; glycine, 2.43 %; alanine, 2.77 %;
and cysteine, 3.71 %.
Plate 10 Lotus leaf petiole peeled and slivered
Nelumbo nucifera 63

The ethanol extract of lotus seeds contained tion of diglycerides, monoglycerides, free
more total fatty acids and total phytosterols than fatty acids and free phytosterol (polar compo-
did the ethanol extract of lotus rhizomes. nents) in lotus seeds (g/kg hexane extract) was
Most of the fatty acids were in the triglyceride determined as diglycerides (1,3-dipalmitoylglyc-
ester form in lotus seeds (Zhou et al. 2013b). erol 0.27 g; 1-palmitoyl-2-linoleoylglycerol
Linoleic acid was the most abundant fatty acid 13.17 g; 1-palmitoyl-3-linoleoylglycerol 32.56 g;
compound in lotus seeds and rhizomes. Fatty 1-oleoyl-2-linolenoylglycerol 4.91 g; 1-oleoyl-3-
acids in lotus seeds existed in the glyceride form linolenoylglycerol 14.44 g), monoglycerides
(59.8 %) and to a lesser extent in their free acid (1-palmitoylglycerol 0.80 g; 2-palmitoylglycerol
form (12.0 %). The concentration of fatty acid 12.64 g; 1-linoleoylglycerol 3.60 g,
and phytosterol in the triglyceride and stearyl- 1-oleoylglycerol 2.06 g; 2-behenoylglycerol
fatty acid ester (non-polar components) in lotus 0.26 g), free fatty acids (myristic (C14:0) 0.31 g,
seeds (g/kg hexane extract) was determined as palmitic (C16:0) 43.26 g, linoleic (C18:2)
myristic (C14:0), 0.86 g; palmitic (C16:0), 26.06 g, linolenic (C18:3) 1.26 g, oleic (cis-
59.46 g; linoleic, (C18:2), 176.4 g; linolenic C18:1) 13.20 g, elaidic (trans-C18:1) 0.74 g,
(C18:3), 6.66 g; oleic (cis-C18:1), 43.19 g; stearic (C18:0) 1.10 g, gondoic (C20:1) 0.45 g,
elaidic (trans-C18:1), 3.87 g; stearic (C18:0), arachidic (C20:0) 0.37 g, behenic (C22:0) 2.24 g,
5.40 g; gondoic (C20:1), 1.47 g; arachidic tricosylic (C23:0) 0.38 g, lignoceric acid (C24:0)
(C20:0), 7.77 g; heneicosylic (C21:0), 1.04 g; 0.85 g, total fatty acids 83.91 g) and free phytos-
behenic (C22:0), 28.94 g; tricosylic (C23:0), terols (campesterol 5.25 g, stigmasterol 0.31 g,
1.64 g; lignoceric acid (C24:0), 8.34 g; total fatty β-sitosterol 56.77 g, β-amyrin 1.61 g, total phy-
acids, 345.97 g; campesterol, 1.55 g; isofucosterol, tosterol 63.94 g). Lotus seed oil was found to be
0.12 g; β-sitosterol, 10.57 g; β-amyrin, 5.34 g; an excellent source of crude oil, 3.52 g/100 g dw
and total phytosterol, 17.58 g. The concentration (Anitha and Arunkumar 2012). The oils had an
of diglycerides, monoglycerides, free fatty acids acid value of 26.66 mg KOH, saponification
and free phytosterol (polar components) in lotus value of 210.43 mg KOH, peroxide value of
seeds (g/kg hexane extract) was determined as 10.60 g/100 g, iodine value of 6.00 and free fatty
diglycerides (1,3-dipalmitoylglycerol, 0.27 g; acids of 3.50 g. Crude fat content of lotus seed oil
1-palmitoyl-2-linoleoylglycerol, 13.17 g; hexane extract was 38.6 %, and fatty acid methyl
1-palmitoyl-3-linoleoylglycerol, 32.56 g; 1-oleoyl- esters found included capric acid methyl ester
2-linolenoylglycerol, 4.91 g; 1-oleoyl-3- (C10:0), lauric acid methyl ester (C12:0), tri-
linolenoylglycerol, 14.44 g), monoglycerides decanoic acid methyl ester (C13:0), palmitic acid
(1-palmitoylglycerol, 0.80 g; 2-palmitoylglycerol, methyl ester (C16:0), pentadecanoic acid methyl
12.64 g; 1-linoleoylglycerol, 3.60 g; ester (C15:0), cis-10-pentadecanoic acid methyl
1-oleoylglycerol, 2.06 g; 2-behenoylglycerol, ester (C15:1), palmitoleic acid methyl ester
0.26 g), free fatty acids (myristic (C14:0), 0.31 g; (16:1), cis-10-heptadecanoic acid methyl ester
palmitic (C16:0), 43.26 g; linoleic (C18:2), (C17:1) and nonadecanoic acid methyl ester
26.06 g; linolenic (C18:3), 1.26 g; oleic (cis- (C19:1).
C18:1), 13.20 g; elaidic (trans-C18:1), 0.74 g; Seed starch grain was oval and contained
stearic (C18:0), 1.10 g; gondoic (C20:1), 0.45 g; higher amylase content starch and gelatinisation
arachidic (C20:0), 0.37 g; behenic (C22:0), temperature than rhizome (Man et al. 2012). Seed
2.24 g; tricosylic (C23:0), 0.38 g; lignoceric acid starches exhibited an A-type X-ray diffraction
(C24:0), 0.85 g; total fatty acids, 83.91 g) and pattern with higher crystalline degree. Seed
free phytosterols (campesterol, 5.25 g; stigmas- starch had lower acid hydrolysis and higher por-
terol, 0.31 g; β-sitosterol, 56.77 g; β-amyrin, cine pancreatic α-amylase hydrolysis than rhi-
1.61 g; total phytosterol, 63.94 g). The concentra- zome starch. A novel lotus plumule polysaccharide
64 Nelumbonaceae

(LPPS) comprising two components F1 and F2 1-(p-methoxybenzyl)-2-methyl-7-isoquinolinol

was purified and characterised (Liao and Lin and d-O-ethylarmepavine were obtained.
2012). The molecular weights of native F1 and Following the isolation of biscoclaurine alka-
F2 were approximately distributed at >2000 and loids, isoliensinine and neferine, from Lien Tze
25.7 kDa, respectively. The total protein and car- Hsin (N. nucifera embryo), from Hong Kong
bohydrate constituent ratios in LPPS, F1 and F2 market, nuciferine and pronuciferine were iso-
were 30.0 % vs. 70.0 %, 30.1 % vs. 69.9 % and lated from the non-phenolic portion and lotusine
96.5 % vs. 3.5 % (w/w), respectively, suggesting from the water-soluble quaternary base portion
that F1 may be a major proteo-polysaccharide (Furukawa 1966a). Also, neferine and liensinine
component and F2 a glycoprotein constituent in were found in lotus seeds from Nepal. Three
LPPS. kinds of phenolic biscoclaurine-type bases were
Nn-9 was a non-crystalline alkaloid isolated obtained from lotus embryo (Lien Tze Hsin) lien-
from lotus embryo (Chen et al. 1962). sinine, isoliensinine and neferine (Furukawa
Bisbenzylisoquinoline alkaloids liensinine (Chao 1966b). A tertiary tetrahydroisoquinoline alka-
et al. 1962; Pan et al. 1962; Hsieh et al. 1964a; loid, methylcorypalline and the biscoclaurine
Guo and Chen 1984) and isoliensinine were iso- base neferine were isolated from lotus embryo
lated from lotus embryo (Tomita et al. 1964; Guo besides the known isoliensinine and lotusine
and Chen 1984). Liensinine was also totally syn- (Yang and Chen 1970a, b).
thesised by Ullmann condensation of 1-(3-bromo- A tetrahydroisoquinoline alkaloid with a sec-
4-benzyloxy-benzyl)-2-methyl-6, ondary base was isolated in crystalline form from
7-dimethoxy-1,2,3,4-tetrahydroisoquinoline and N. nucifera embryo and identified with
1-(4-benzyloxy-benzyl)-2-methyl-6-methoxy-7- 1-(p-hydroxybenzyl)-6, 7-dihydroxy-1, 2, 3,
hydroxy-1,2,3,4-tetrahydroisoquinoline (Hsieh 4-tetrahydroisoquinoline (demethylcoclaurine)
et al. 1964b). Pronuciferin is a benzylisoquino- (Koshiyama et al. 1970). Three kinds of phenolic
line with para-cyclohexadienone grouping biscoclaurine-type bases liensinine, isoliensinine
(Bernauer 1963, 1964); the noraporphine alka- and neferine plus O-dimethylliensinine were
loid anonaine, besides the aporphines nuciferine, obtained from lotus embryo (Lien Tze Hsin) of
roemerine and 5-methoxy-6-hydroxy-aporphine, lotus (Furukawa 1971). The structure of nor-
was isolated from lotus cotyledons (Bernauer nuciferine, an aporphine-type tertiary phenolic
1964). Isoliensinine, a new phenolic base isolated from the domestic lotus, was proved
biscoclaurine-type alkaloid, was isolated from to be correct (Tomita et al. 1971). From lotus
Formosan “Lien Tze Hsin,” lotus seed embryo receptacle, four alkaloids, nuciferine,
(Tomita et al. 1965). A new, water-soluble, qua- N-nornuciferine, oxoushinsunine and
ternary base alkaloid named lotusine was isolated N-norarmepavine, were isolated (Yang et al.
from a Formosan lotus embryo besides the earlier 1972). Isoliensinine, neferine, (±)-armepavine
reported isoliensinine (Furukawa et al. 1965). A and 4′-methtyl-N-methylcoclaurine were iso-
new base, neferine as well as isoliensinine, were lated from lotus embryo (Nishibe et al. 1986).
isolated from “Lien Tze Hsin” (Nelumbo nucifera Six alkaloids lotusine, nuciferine, pronuciferine,
embryo), from Hong Kong market (Furukawa liensinine, isoliensinine and neferine were iso-
1965a). Liensinine was found to be the main lated from lotus green seed embryo (Wang et al.
alkaloid in Japanese lotus embryo, besides a 1991). Liensinine was extracted from lotus embryo
small amount of neferine (Furukawa 1965b). by impregnating and refluxing, yielding content of
O-Methylneferine (styphnate) was found to be 0.853 % and 0.939 % and with average recovery of
completely identical with the synthesised O, 97.9 % and 100.9 %, respectively (Hu et al. 1993).
O-dimethylisoliensinine, reported previously. The pronucerine monomer was isolated from
When O-methylneferine was subjected to the lotus seed (Xiao et al. 2007). Using the first sol-
cleavage reaction by metallic sodium in liquid vent system of light petroleum–ethyl acetate–tet-
ammonia, d-1, 2, 3, 4-tetrahydro-6-methoxy- rachloromethane–chloroform–methanol–water
Nelumbo nucifera 65

solvent system, 1102 mg of the crude lotus glucopyranoside and apigenin-6-O-α-l-

embryo was purified yielding 350 mg neferine, arabofuranosyl-8-O-β-d-glucopyranoside, and
100 mg isoliensinine and 95 mg liensinine with from seeds apigenin-6-O-α-l-arabofuranosyl-8-
over 95 % purity (Wu et al. 2004). While using O-β-d-glucopyranoside and unknown compound
the second solvent system ethyl acetate–tetra- (Zhou et al. 2013a).
chloromethane–methanol–water, 5850 mg of the The flavonoid content of lotus seed embryos
crude alkaloid was purified yielding 2545 mg was high, 730.95 mg 100/g DW (dry weight),
neferine, 698 mg isoliensinine and 650 mg lien- seed coats had much less total flavonoid
sinine with over 97 % purity. Three bisbenzyl- (<40 mg/100 g FW) and seed kernels had no
isoquinoline alkaloids (liensinine, isoliensinine flavonoids (Chen et al. 2012b). Seed coats con-
and neferine) were isolated from lotus embryo tained myricetin 3-O-glucoside; quercetin
(Chen et al. 2007). The mass fractions of liensi- 3-O-arabinopyranosyl-(1 → 2)-galactopyrano-
nine, isoliensinine and neferine in the crude side; myricetin 3-O-glucuronide; quercetin
extract and the phenolic alkaloid sample of lotus 3-O-rhamnopyranosyl-(1 → 6)-glucopyranoside
embryo were 16.5 and 228.6 mg/g for liensinine, (rutin); quercetin 3-O-galactoside (hyperoside),
45.7 and 640.7 mg/g for isoliensinine and 59.7 quercetin 3-O-glucoside (isoquercitrin); quercetin
and 58.8 mg/g for neferine, respectively. 3-O-glucuronide; isorhamnetin 3-O-rutinoside;
Neferine, a bisbenzylisoquinoline alkaloid, is kaempferol 3-O-glucoside (astragalin); syringetin
extracted (isolated) from the green seed embryo 3-O-glucoside; isorhamnetin 3-O-glucoside and
(Chen et al. 2008). From 200 mg of crude extract, kaempferol 3-O-glucuronide. Fruit coats con-
18.4 mg of liensinine, 19.6 mg of isoliensinine tained similar flavonoids plus kaempferol
and 58.4 mg of neferine were obtained with the 3-O-galactoside; diosmetin 7-O-hexose; isorham-
purity of 96.8, 95.9, and 98.6 %, respectively netin 3-O-glucuronide; quercetin and diosmetin.
from lotus seed embryo (Liu et al. 2009). In a Four known alkyl 4-hydroxybenzoates, i.e. methyl
separate study, 200 mg of crude lotus embryo 4-hydroxybenzoate; ethyl 4-hydroxybenzoate;
extract, 33 mg of liensinine, 42 mg of isoliensi- propyl 4-hydroxybenzoate and butyl
nine and 67 mg of neferine were obtained all 4-hydroxybenzoate, were isolated from lotus
with purities over 98 % (Duanmu et al. 2010). A seeds (Youn et al. 2010). Four aglycones of Fr2
2.5 g crude alkaloid extract from lotus seed fraction from lotus seed epicarp (seed coat) were
yielded 151 mg of liensinine, 118 mg of isolien- identified as myricetin, quercetin, kaempferol and
sinine and 572 mg of neferine with purities of isorhamnetin (Kredy et al. 2010). The results indi-
93.0, 95.1 and 97.0 %, respectively (Wang et al. cated that the quercetin content in Fr2 (10.2 mg/g
2010). From the embryos of lotus seeds, three of dry fraction) was higher than isorhamnetin
bisbenzylisoquinoline alkaloids, nelumboferine (6.2 mg/g of dry fraction), followed by kaemp-
and nelumborines A and B, were isolated along ferol (2.8 mg/g of dry fraction) and myricetin
with four known compounds, neferine, liensi- (0.4 mg/g of dry fraction). Six glycosylated
nine, isoliensinine and anisic acid (Itoh et al. flavonols were found in Fr2 fraction; they were
2011). Bisbenzylisoquinoline alkaloid, nelum- myricetin monohexosides (may be myricetin
boferine which was recently isolated from the 3-O-galactoside or myricetin 3-O-glucoside);
embryo of Nelumbo nucifera and stereoisomers quercetin diglycoside, confirmed as rutin
of neferine, a major alkaloid of the embryo of N. (quercetin 3-O-rutionoside); quercetin monohexo-
nucifera, were stereoselectively synthesised side (the hexose could be glucoside or galactoside);
(Nishimura et al. 2013). Ten major anti-inflam- kaempferol-monodeoxyhexoside-monohexoside,
matory compounds were identified from lotus possibly kaempferol 3-O-rutinoside; isorhamnetin-
plumule, i.e. higenamine, lotusine, 4′-methylco- monodeoxyhexoside-monohexoside, possibly
claurine, isoliensinine, liensinine, neferine, isorhamnetin 3-O-rutinoside; and an isorhamnetin
armepavine, 4′-methyl-N-methylcoclaurine, monohexoside (the hexose may be glucoside or
apigenin-6- O -α- l - glucopyanosyl-8- O -β- d - galactoside) (Kredy et al. 2010).
66 Nelumbonaceae

The procyanidins of lotus seedpod were heat-induced annexin, NnANN1, was identified
extracted with aqueous acetone with a purity of from lotus embryonic axes that was found to play
>98 % (Lin et al. 2005). The main molecular an important role in seed thermotolerance and
weight distribution of procyanidins ranged from germination vigour. Compared to the wild-type
291 to 1155, with M + H peak values of 291.1, seeds, transgenic seeds ectopically expressing
579.2, 731.2, 867.2, 1019.4 and 1155.3, respec- NnANN1 exhibited improved resistance to accel-
tively, indicating the extract contained mono- erated ageing treatment. Also transgenic seeds
mers, dimers and tetramers of procyanidins, in showed enhanced peroxidase activities, accom-
which the amounts of dimers were greatest, and panied with reduced lipid peroxidation and
catechin and epicatechin were the base units. The reduced ROS release levels compared to the
procyanidins of lotus seedpod were extracted wild-type seeds. Studies by Shen-Miller et al.
with aqueous acetone with a purity of >98 % (Lin (2013) showed that in proteins of N. nucifera
et al. 2005). The main molecular weight distribu- fruit that were able to withstand heating, 31 % of
tion of procyanidins ranged from 291 to 1155, which remained soluble in the 110 °C-treated
with M + H peak values of 291.1, 579.2, 731.2, embryo-axis of a 549-yr-old fruit and 76 %
867.2, 1019.4 and 1155.3, respectively, indicat- retained fluidity in its cotyledons. At 110 °C for
ing the extract contained monomers, dimers and 10 min, 22.6 μg/mg of thermal proteins were
tetramers of procyanidins, in which the amounts recovered from the embryo-axis of a 549-year-
of dimers were greatest, and catechin and epicat- old lotus seed versus 7.1 μg/mg from modern
echin were the base units. Five flavonol glyco- seeds; 11.1 μg/mg were recovered from the coty-
sides, including hyperoside, isoquercitrin, ledons of a 549-year-old seed versus 21.9 μg/mg
quercetin-3-O-ß-d-glucuronide, isorhamnetin-3- from the cotyledons of modern seeds. The amino-
O-ß-d-galactoside and syringetin-3-O-ß-d-- acid sequences of 11 “thermal proteins” (soluble
glucoside, were determined in lotus receptacle at 100 °C) of modern Nelumbo embryo-axes and
with recoveries in the range of 98.31–100.32 % cotyledons were identified as dehydrin (gluta-
(Wu et al. 2013). The oligomeric and polymeric mine, lysine), 55 % total amino acid; HSP80
fraction of the 60 % aqueous methanol lotus seed (lysine/glutamine, asparagine), 38 %; CPN60
pod extract had a mean degree of polymerisation (lysine, glutamine), 33 %; vicilin (glutamine,
of 3.2 and 15.4, respectively, and consisted of lysine, arginine/asparagine), 30 %; EF-1α (lysine,
(+)-catechin (m/z 289), gallocatechin or epigal- asparagine, glutamine), 29 %; met-synthase (glu-
locatechin (m/z 305), quercetin glycoside (m/z tamine, lysine, asparagine), 36 %; 1-CysPRX
463), quercetin glucuronide (m/z 477), procyani- (lysine, asparagine), 26 %; ENO1 (lysine, gluta-
din dimers (m/z 577.1), proanthocyanidin dimer mine), 24 %; CPN20 (lysine, glutamine, aspara-
gallate (m/z 593.3), prodelphinidin dimers (m/z gine), 22 %; PIMT (glutamine, lysine,
609.1), procyanidin trimers (m/z 865.1), etc asparagine), 21 %; and CuZn-SOD (asparagine,
(Xiao et al. 2012). Quercetin glucuronide was lysine, glutamine), 18 %. The 11 Nelumbo
further purified and identified as quercetin thermal proteins studied could be grouped into
-3-O-β-glucuronide. A flavonoid quercetin [2- four functionally defined categories: antioxidant
(3,4-dihydroxy)-3, 5, 7-trihydroxy-4H-1-benzo- and membrane maintenance (CuZn-SOD,1-
pyran-4-one] was isolated from lotus dried recep- CysPRX, dehydrin); chaperonins, chaperone and
tacle (Ishida et al. 1988). Five flavonol glycosides, stress proteins (Cpn20, Cpn60, HSP80, EF-1α);
namely, hyperoside, isoquercitrin, quercetin- defence, food and repair (vicilin, Met-Synthase,
3-O-β-d-glucuronide, isorhamnetin-3-O-β-d- PIMT); and anaerobic glycolytic protein
galactoside and syringetin-3-O-β-d-glucoside, (Enolase1 (ENO1)). In addition, 30 other heat-
were isolated from lotus receptacle (Wu et al. stable proteins of N. nucifera cv. China Antique
2012). were identified: actin 12, aldose reductase, aller-
Lotus seeds were found to contain annexins, gen Ara h1 clone, nep1 aspartic proteinase
multifunctional proteins (Chu et al. 2012). A nepenthesin 1, chaperone protein yajL, embry-
Nelumbo nucifera 67

onic protein DC-8, pistil-specific extensin-like drates, 17.23 g; total dietary fibre, 4.9 g; Ca,
protein, FBA fructose biphosphate aldolase 45 mg; Fe, 1.16 g; Mg, 23 mg; P, 100 mg; K,
(cytosol enzyme), 11S globulin β, FA02 13S 556 mg; Na, 40 mg; Zn, 0.39 mg; Cu, 0.257 mg;
globulin seed storage protein 1, GluA1 glutelin Mn, 0.261 mg; Se, 0.7 μg; vitamin C, 44 mg;
type A1, GluB1 glutelin type B1, GAPC thiamine, 0.160 mg; riboflavin, 0.220 mg; niacin,
glyceraldehye-3-phosphate dehydrogenase 0.400 mg; pantothenic acid, 0.377 mg; vitamin
(cytosol), HSP18.2 18.2 kDa Class 1 heat shock B-6, 0.258 mg; total folate, 13 μg; total saturated
protein and HSP70; late embryonic abundant fatty acids, 0.030 g; 16:0 (palmitic acid), 0.028 g;
(LEA) proteins Dc3, Dc 8 and Dc 34; and 18:0 (stearic acid), 0.001 g; total monounsatu-
Legumin A (LEGA), Legumin B (LEGB), low rated fatty acids, 0.020 g; 16:1 undifferentiated
temperature-induced 65 kDA (LT65) protein, (palmitoleic acid), 0.002 g; 18:1 undifferentiated
malate dehydrogenase (MDH), 5-methyltetrahyd (oleic acid), 0.014 g; 20:1 (eicosenoic acid),
ropteroyl-triglutamate-homocysteine methyl- 0.002 g; total polyunsaturated fatty acids,
transferase (metE), early nodulin-like protein 0.020 g; 18:2 undifferentiated (linoleic acid),
1(Atg25060), protein notum homolog (NOTUM), 0.014 g; 18:3 undifferentiated (linolenic acid),
pectiesterase 2 (PECS-2.1), PCKR 1 peptidyl- 0.02706 g; tryptophan, 0.020 g; threonine,
prolyl cis-trans isomerise, peroxidase 12 0.051 g; isoleucine, 0.054 g; leucine, 0.069 g;
(PER12), phosphoglycerate kinase (cytosol) and lysine, 0.094 g; methionine, 0.022 g; cystine,
tricohyalin and triose-phosphate isomerase 0.022 g; phenylalanine, 0.047 g; tyrosine,
(TCHH). 0.029 g; valine, 0.055 g; arginine, 0.088 g; histi-
The main constituents in the essential oils dine, 0.038 g; alanine, 0.054 g; aspartic acid,
from the various rhizome and seed samples were 0.369 g; glutamic acid, 0.139 g; glycine, 0.156 g;
l-(+)-ascorbic acid 2,6-dihexadecanoate proline, 0.136 g; and serine, 0.060 g (USDA-
(0–33.5 %), trans-phytol (5.1–24.1 %), hexahy- ARS 2014).
drofarnesyl acetone (5.6–15.3 %), pentadecyl The nutrient composition of the rhizome per
acrylate (2.2–12.4 %), geranyl acetone (1.9– 100 g edible portion (CINE 2007) was reported
8.0 %) and β-ionone (0–8.0 %) (Huang et al. as follows: moisture, 80.5 g; energy, 75 kcal; pro-
2010a). The rhizome lotus and seed lotus sam- tein, 1.9 g; fat, 0.2 g; carbohydrate, 16.4 g; fibre,
ples were clustered into separate groups by hier- 1.2 g; ash, 1 g; vitamin A, 3 μg RE (1.5 μg RAE);
archical cluster analysis according to the total carotene, 20 μg; vitamin C, 44 mg; Zn,
composition of the corresponding essential oils. 0.23 mg; Fe, 1.4 mg; and Ca, 39 mg. Another
No significant relationship was found between study reported that the rhizomes have thiamine,
essential oil composition and geographical distri- 0.16 mg; riboflavin, 0.22 mg; niacin, 0.4 mg;
bution of the 11 populations. vitamin C, 44 mg; calcium, 45 mg; phosphorus,
Young chloroplasts were found in shoots of 100 mg; iron, 1.6 mg; potassium, 730 mg;
mature dry lotus seeds, and they gave rise to sodium, 40 mg; magnesium, 25 mg; zinc, 0.2 mg;
mature chloroplasts during germination, even in protein, 2.6 g; carbohydrate, 14.7 g; and fat, 0.1 g
darkness (Ushimaru et al. 2003). These shoots (Howard et al, 1962). Korean white lotus culti-
contained chlorophyll and chlorophyll-binding vars (Inchisa, Muan, Garam and Chungyang) and
proteins CP1 and LHCP. rhizomes were reported to possess high amounts
of bioactive compounds: total phenols, between
7.95 and 4.21 mg of gallic acid equivalents
Rhizome Nutrients/Phytochemicals (GAE)/g dry weight (DW); ascorbic acid,
between 15.8 and 22.3 mg of ascorbic acid/g
The nutrient composition of raw rhizome per DW; and amino acids, between 15.05 % and
100 g edible portion was reported as follows: 16.62 0 % DW (Park et al. 2009). Tryptophan was
water, 79.10 g; energy, 74 kcal (311 kJ); protein, isolated from the aqueous extract of lotus rhi-
2.60 g; total lipid, 0.10 g; ash, 0.97 g; carbohy- zome (Jiang et al. 2010).
68 Nelumbonaceae

The ethanol extract of lotus rhizomes con- respectively, while least gelation concentration,
tained less total fatty acids and total phytosterols foaming volume increase, foaming stability,
than did the ethanol extract of lotus seeds (Zhao emulsifying capacity and emulsion stability were
et al. 2013a). The concentration of fatty acid and 18.0 %, 5.23 %, 4.97 %, 48.93 % and 96.43 %,
phytosterol in the triglyceride and stearyl-fatty respectively. The lotus rhizome was found to be a
acid ester (non-polar components) in lotus rhi- poor source of crude oil (2.68 g/100 g dry weight)
zomes (g/kg hexane extract) was determined as with the following physicochemical properties:
follows: palmitic (C16:0), 43.64 g; linoleic crude fat (2.68 g/100 g DW), acid value (16.66 mg
(C18:2), 53.68 g; linolenic (C18:3), 7.91 g: KOH), saponification value (110.43 mg KOH)
oleic (cis-C18:1), 14.27 g: elaidic (trans- and unsaponifiable matter (0.026 g/100 g oil).
C18:1), 1.17 g; stearic (C18:0), 8.34 g; ara- The amylose content was the highest
chidic (C20:0), 0.43 g; behenic (C22:0), 1.56 g; (30.61 %) in lotus starch (Zhiong et al. 2007).
and total fatty acids, 131.01 g. The concentra- The average particle size (diameter) was 50.27,
tion of diglycerides, monoglycerides, free fatty 24.08 and 38.97 μm for lotus, kudzu and corn
acids and free phytosterol (polar components) starches, respectively. Lotus starch exhibited a
in lotus rhizomes (g/kg hexane extract) was B-type X-ray diffraction pattern and kudzu starch
determined as follows: diglycerides (0.27 g exhibited a C-type pattern. Kudzu starch was
1-palmitoyl-2-linoleoylglycerol and 0.45 g characterised by a maximum viscosity immedi-
1-palmitoyl-3-linoleoylglycerol), monoglycer- ately followed by a sharp decrease in viscosity,
ides (8.13 g 2-palmitoylglycerol and 7.11 g while the lotus starch was characterised by a pla-
1-linoleoylglycerol), free fatty acids (myristic teau when the maximum viscosity was reached.
(C14:0), 0.11 g; palmitic (C16:0), 6.80 g; linoleic Lotus rhizome starches had small oval gran-
(C18:2), 5.21 g; linolenic (C18:3), 1.51 g; oleic ules and large elongated granules (Man et al.
(cis-C18:1), 1.51 g; elaidic (trans-C18:1), 0.15 g; 2012). Rhizome starches had higher swelling
stearic (C18:0), 0.41 g; gondoic (C20:1), 0.02 g; power, lower amylase content and gelatinisation
arachidic (C20:0), 0.09 g; heneicosylic (C21:0), temperature than seed starch. Rhizome starches
0.02 g; behenic (C22:0), 0.34 g; tricosylic showed a C-type pattern which changed from C
(C23:0), 0.08 g; lignoceric acid (C24:0), 0.45 g; to A type with gradually increasing crystalline
and total fatty acids, 83.9116.02 g) and free phy- degree during acid hydrolysis. The degree of
tosterols (campesterol, 4.51 g; isofucosterol, order in starch external region was higher in rhi-
0.65 g; β-sitosterol, 76.27 g; β-amyrin, 7.34 g; zome than in seed. The external region structure
α-amyrin, 7.78 g; lanosterol, 12.68 g; and total of rhizome starches became more ordered during
phytosterol, 109.22 g). acid hydrolysis. Rhizome starches had higher
The proximate composition of lotus rhizome rate of acid hydrolysis and lower rate of porcine
flour (g/100 g flour) was determined as ash pancreatic α-amylase hydrolysis than seed
(1.10 g), the total nitrogen (1.36 g), total protein starches. Starch in rhizome of lotus cultivar
(8.48 g), water-soluble proteins (1.23 g), salt- Meirenhong exhibited C-type X-ray diffraction
soluble proteins (5.73 g), total salt-soluble pro- pattern, while starch in rhizome of cultivar
teins (6.064 g), total sugar (19.08 g), reducing Wawalian showed A-type pattern (Yu et al.
sugars (0.168 g), non-reducing sugars (18.87 g) 2013a). Starch granules in cv. Meirenhong
and free amino acids (0.78 g) (Shad et al. 2011). showed oval-shaped granules, while starch gran-
The temperature-dependent behaviour of solubil- ules in cv. Wawalian were elongated and oval in
ity and swelling capacity of the rhizome flour shape with relatively large size. Gelatinisation
showed a linear increase (1.2–13.84 %) in solu- temperatures of starch in Meirenhong and
bility but an exponential increase in swelling Wawalian were 330.5 and 342.4 K, respectively,
capacity with a gradual increase in the tempera- and the gelatinisation temperature range of
ture (40–100 °C). Water absorption and oil hold- Meirenhong was significantly wider than that of
ing capacities were found to be 2.56 and 2.03 %, Wawalian. Starch in rhizome of cv. Meirenhong
Nelumbo nucifera 69

showed lower pasting temperature, lower hot and extracts yielded alkaloids, steroids, reducing sug-
cool viscosities, lower setback and higher peak ars and saponins (Mukherjee et al. 1998). The
viscosity and breakdown than those of cv. detection and quantification limitations of cate-
Wawalian in rapid viscosity analyser pasting pro- chin were 23 ng and 50 ng, respectively, in lotus
files at 6 % starch concentration. Amylose mole- rhizome (Yan et al. 2009). The catechin level was
cules in lotus rhizome starch were large, with 0.0025 % in the lotus rhizome and 0.011 % in the
number-average and weight-average degrees of knot of the lotus rhizome. (±)-Gallocatechin and
polymerisation (DPs) of 4170 and 8040, respec- (–) catechin plus LB2 was a polysaccharide-
tively, number-average chain length (͞͞CLn) of 540 protein complex with a molecular mass of
and apparent DP distribution range of 520– 18.8 kDa isolated from lotus rhizome (Jiang et al.
42,000 (Suzuki et al. 1992). The ͞CLn of amylo- 2011). LB2 was identified as a polysaccharide
pectin was 22.3, and its distribution by sulphate containing α/β-pyranose and α-furanose.
gel-exclusion HPLC showed three peaks, at DP It was composed of mannose, rhamnose, glucose,
14, 42 and 1900, and a shoulder at DP 63. The galactose and xylose with a molar ratio 2:8:7:8:1.
amylose content was 15.9 % determined by The percentage of recovery of betulinic acid
iodine affinity and 17.4 % determined by blue from lotus rhizome by HPTLC was found to be
value. The onset of gelatinisation determined by 98.36 % (Mukherjee et al. 2010c). The limit of
photopastegraphy was at 58.5 °C. The amylo- detection and limit of quantification of betulinic
gram of the starch resembled that of tapioca acid were detected to be 0.4 and 2.30 μg per spot.
starch. The retrogradation tendency of lotus From lotus rhizome, a new ursane triterpenoid
starch was slower than that of potato, lily and ester, urs-12-en-3β-O-9E,12E-octadecadienoate,
kudzu starches but faster than that of tapioca was identified along with the isolation of seven
starch. During gelatinisation of lotus rhizome known compounds (palmitic acid, linoleic acid,
starch, the crystallinity changed from C type to A 9E,12E,15E-octadecatrienoic acid, α-amyrin,
type via CA type and finally became amorphous β-sitosterol, betulinic acid and β-sitosterol-3-O-
structure (Cai et al. 2014). The amylose content, glucoside) (Chaudhuri and Singh 2009) and a
crystal degree, helix content, ratio of new triterpenoid, 2α,24-diacetoxy-3β-
1045/1022 cm−1 and peak intensity of crystalline hydroxyolean-12-en-28-oic acid, along with four
lamellae of gelatinising starch significantly known compounds (hyptatic acid-A (2α,3β,24-
decreased after 70 °C. The amorphous content trihydroxyolean-12-en-28-oic acid), maslinic
and ratio of 1022/995 cm−1 increased after acid (2α,3β-dihydroxyoleane-12-ene-28-oic
70 °C. The study elucidated that B-type allo- acid), betulin and lupeol) (Chaudhuri and Singh
morph was mainly arranged in the distal region 2013).
of eccentric hilum, A-type allomorph was mainly Fifteen compounds were isolated from the
located in the periphery of hilum end, and the total alkaloid extract of lotus stem (rhizome) and
centre of the starch granule was a mixed distribu- identified as asimilobine, isococlaurine,
tion of A- and B-type allomorphs. Lotus seed- N-acetylnorarmepavine, crykonisine, velucryp-
resistant starch (LRS), a type of retrograded tine, pycnarrhine, liriodenine, nuciferine, nor-
starch, is commonly known as resistant starch nuciferine, armepavine, N-methylasimilobine,
type 3 (RS3) (Zhang et al. 2014). Compared with coclaurine, N-norarmepavine (13),
native starch and high amylose maize starch N-methylcoclaurine and lysicamine (Duan et al.
(HAMS), LRS lacked the polarisation cross, and 2013). Astragalin, rutin, isoquercetin, nuciferine,
the irregularly shaped LRS granules had a dauricine, isoliensinine and neferine were identi-
rougher surface, B-type crystal structure and fied in lotus rhizomes (Zhao et al. 2014).
greater level of molecular order. High-quality RNA was recovered from five
The petroleum ether extract of lotus rhizomes different tissues of lotus as assessed by polyphe-
yielded steroids; the chloroform extract yielded nol oxidase (PPO) gene expression profiles
alkaloids and steroids; the methanol and water (Zhang et al. 2013b). The relative expression lev-
70 Nelumbonaceae

els of the PPO gene based on RTPCR in five tis- zome (Tomita et al. 1961c). Roemerine and
sues of lotus were rhizome buds (2.66), young nuciferine were isolated as the tertiary, non-
leaves (2.42), fresh cut rhizome (2.02), petals phenolic bases, and a new base, nornuciferine, as
(1.80) and petiole (1.65), using housekeeping the tertiary phenolic base, from the leaves.
gene β-actin as an internal control. The ethanol Roemerine and nornuciferine were isolated from
extracts of lotus seeds from Vietnam (Ho Chi the petiole but not from nuciferine. The results
Minh City), raw rhizomes from Korea (Siheung) indicated that the main alkaloids of lotus are all
and dried rhizomes from Japan (Nigata) had the tertiary bases of the aporphine type. Roemerine,
greatest specific flavonoid content (Zhao et al. nuciferine and nornuciferine, a new aporphine-
2014). The ethanol extracts of seeds from China type tertiary phenolic base, were isolated from
(Hubei), raw rhizomes from Japan (Nigata) and the leaves and petioles of domestic lotus (Tomita
dried rhizomes from Korea (Siheung) had the et al. 1961a). Besides the isolation of phenolic
greatest specific alkaloid content. bases, nuciferine, roemerine and nornuciferine,
another base, dl-armepavine, was isolated from
the leaves (Tomita et al. 1961b). Aporphine-type
Leaf Nutrients/Phytochemicals alkaloid bases, roemerine, and nornuciferine,
and benzyl-tetrahydro-isoquinoline-type base,
Proximate nutrient composition of lotus leaf (g armepavine, from the leaves, and roemerine and
DW/100 g) was reported as moisture, 1.65 g; nornuciferine from the leaf petiole were isolated
crude protein, 14.47 g; crude lipid, 2.29 g; crude from an ancient lotus cultivar Ohga-hasu (Tomita
ash, 9.36 g; carbohydrate, 10.23 g; chemicals and Frurukawa 1962). Alkaloids contained in the
(mg DW)/100 g K, 1345.25 mg; Ca, 1069.51 mg; so-called primitive lotus ‘gen-shi-hasu’ were
P, 293.27 mg; Mg, 261.48 mg; Na, 12.4 mg; Fe, similar to domestic lotus: roemerine, nuciferine,
4.32 mg; Zn, 1.76 mg; Cu, 0.26 mg; sucrose, nornuciferine and dl-armepavine (Kunitomo
1646.56 mg; fructose, 210.70 mg; glucose, et al. 1964). Cleavage reaction of nuciferine and
110.14 mg; vitamin A, 714.61 mg; vitamin C, nornuciferine by metallic sodium in liquid
679.59 mg; vitamin B1, 4.10 mg; and vitamin ammonia afforded 2-methoxyaporphine and
B2, 3.67 mg (Ha et al. 2010). 2-hyrdoxyaporphine, respectively (Kunimoto
Scanning electron microscopy and chemical and Kamimura 1964). Tertiary alkaloids
analysis of the chloroform extract of lotus leaves anonaine, pronuciferine, N-nornuciferine, liri-
showed that the wax was composed of a mixture odenine and d-methylcoclaurine were isolated
of aliphatic compounds, principally nonacosanol from the leaves besides the known roemerine,
and nonacosanediols (Koch et al. 2006). Analysis nuciferine, O-nornuciferine and dl-armepavine
of gas chromatography spectra of lotus leaf (Kunitomo et al. 1970). The structure of nor-
waxes showed a much lower proportion of the nuciferine, an aporphine-type tertiary phenolic
secondary alcohol nonacosan-10-ol (16.2 % by base isolated from the domestic lotus, was proven to
weight) compared with nonacosanediols be dl-1-methoxy-2-hydroxyaporphine (Kunitomo
(64.7 %). Gas chromatographic analysis of the et al. 1971). Four new alkaloids, dehydroro-
extracted leaf waxes revealed nonacosan-10-ol emerine, dehydronuciferine, dehydroanonaine
(16.2 %), triacontan-7-ol (2.4 %), nonacosane-4, and N-methylisococlaurine, were isolated from
10-diol (18.6 %), nonacosine-5, 10-diol (34.1 %), N. nucifera leaves, besides the known roemerine,
nonacosane-10, 13-diol (12.0 %), hentriacon- nuciferine, anonaine, pronuciferine,
tane- 12, 15-diol (1.8 %), tritriacontane-9, 10-diol N-nornuciferine, nornuciferine, amepavine and
(0.7 %) and octadecanoic acid (0.7 %). N-methylcoclaurine (Kunitomo et al. 1973).
The first alkaloid isolated from lotus leaves Asimilobine and lirinidine, serotoninergic recep-
was nuciferine (Arthur and Cheung 1959). tor antagonists, were isolated from lotus leaves
Alkaloids were found in lotus leaves, petiole and (Shoji et al. 1987). Four alkaloids, i.e., 2-hydroxy-
cotyledons in the fruit but not in the edible rhi- 1-methoxyaporphine, pronuciferine, nuciferine
Nelumbo nucifera 71

and roemerine, were identified in Nelumbo nuciferine and nornuciferine, was isolated from
nucifera and its alkaloid fraction by HPLC (Wang Nelumbo nucifera leaves (Yang et al. 2012). The
et al. 2008). The linear ranges of 2-hydroxy-1- following major alkaloids (−)-nuciferine,
methoxyaporphine, pronuciferine, nuciferine and (−)-nornuciferine, (−)-caaverine, (−)-armepavine,
roemerine were 0.110–0.658 μg (R2 = 0.9995), (+)-norarmepavine, (+)-isoliensinine and
0.0210–0.126 μg (R2 = 0.9995), 0.103–0.618 μg (+)-pronuciferine were found in Nelumbo
(R2 = 0.9998) and 0.085 6–0.514 μg (R2 = 0.9995), nucifera leaves (Do et al. 2013). In all samples,
with the average recoveries of 101.5 %, 99.14 %, (−)-nuciferine (0.34–0.63 %), (−)-armepavine
99.21 % and 98.41 % for the alkaloid fraction of (0.13–0.20 %) and (+)-isoliensinine (0.06–
N. nucifera and 99.53 %, 100.5 %, 97.51 % 0.19 %) were the most dominant alkaloids. The
and 100.1 % for N. nucifera, respectively. total alkaloid content varied from 0.72 to 1.41 %.
(+)-1(R)-Coclaurine, (–)-1(S)-norcoclaurine, From the methanolic extracts of lotus flower buds
liensinine, neferine, isoliensinine and nuciferine and leaves, a new alkaloid, N-methylasimilobine
together with quercetin 3-O-β-d-glucuronide (4) N-oxide, was isolated together with 11 benzyliso-
were isolated from lotus leaves (Kashiwada et al. quinoline alkaloids (Nakamura et al. 2013).
2005). N-Nornuciferine, O-nornuciferine, nucif- 2-Hydroxy-1-methoxyaporphin (2H1M), an
erine and roemerine were identified in lotus alkaloid, was isolated from lotus leaves (Chen
leaves (Luo et al. 2005). From 4.0 g of the crude et al. 2012c). Major anti-inflammatory com-
lotus leaf extract, 120 mg N-nornuciferine, pounds identified from lotus leaves were nucifer-
1020 mg nuciferine and 96 mg roemerine were ine, O-nornuciferine, hirsutrin, hyperin,
obtained each with a purity of over 98 % (Zheng luteoloside and roemerine (Zhou et al. 2013a).
et al. 2010). Three alkaloids N-nornuciferine, From lotus leaves, liriodenine, lysicamine,
O-nornuciferine and nuciferine were isolated (−)-anonaine, (−)-asimilobine, (−)-caaverine,
from lotus leaf (Ma et al. 2010). 500 mg of crude (−)- N -methylasimilobine, (−)-nuciferine,
lotus leaf extract furnished 7.4 mg of (−)-nornuciferine, (−)-roemerine, cepharadione
N-demethylarmepavine, 45.3 mg of nuciferine B and 7-hydroxydehydronuciferine were isolated
and 26.6 mg of roemerine with purities of 90 %, (Lin et al. 2014). 100 mg of crude lotus leaf
92 % and 96 %, respectively (Xu et al. 2011b). extracts yielded 6.3 mg of 2-hydroxy-1-
The compounds isolated from lotus leaves methoxyaporphine (95.1 % purity), 1.1 mg of
included seven aporphines ((–)-nuciferine, pronuciferine (96.8 % purity), 8.5 mg of nucifer-
(–)-nornuciferine, (–)-asimilobine,(–)-N- ine (98.9 % purity) and 2.7 mg of roemerine
methylasimilobine, (–)-caaverine, (–)-anonaine (97.4 %) (Ma et al. 2014).
(6) and (–)-roemerine), two oxoaporphines (lysi- Quercetin, myricetin, kaempferol
camine and liriodenine), one dioxoaporphine 3-O-glucoside, quercetin 3-O-glucoside and
(cepharadione B), one dehydroaporphine luteolin 7-O-glucoside were isolated from aer-
(–7-hydroxydehydronuciferine), two steroids ial parts of N. nucifera (Wassel et al. 1996).
(β-sitostenone and stigmasta-4,22-dien-3-one) From lotus leaf extract, the following com-
and two chlorophyll (pheophytin-a and aristo- pounds were isolated: a new megastigmane,
phyll-C) (Wang et al. 2011). (–)-caaverine, ceph- nelumnucifoside, together with 12 megastig-
aradione B and 7-hydroxydehydronuciferine manes ((E)-3-hydroxymegastigm-7-en-9-one,
were reported for the first time in lotus. (−) –boscialin, (+) – dehydrovomifoliol, vomi-
Liriodenine in lotus leaves was extracted and foliol, 3-oxo-retro-α-ionol I, byzantionoside A,
quantified using ethanol and water-soluble sol- 5,6-epoxy-3- hydroxy-7-megastigmen-9-one,
vents; liriodenine content obtained was 4.8 mg% annuionone D, icariside B2, grasshopper
by both methods (Mongkolrat et al. 2012). ketone, 3S, 5R-dihydroxy-6S, 7-megastigma-
N-methylasimilobine, a newly found potent dien-9-one and (+)-epiloliolide); a new eudes-
acetylcholinesterase (AChE) inhibitor, along mane sesquiterpene, eudesnelumboside; eight
with two weakly active aporphine alkaloids, alkaloids ((6R, 6aR)-roemerine-Nβ-oxide,
72 Nelumbonaceae

liriodenine, pronuciferine, oleracein E, trans- quercetin-3-O-glucuronide, quercetin-3-O-galac-

N-coumaroyltyramine, cis-N-coumaroyltyramine, topyranoside, kaempferol-3-O-glucopyranoside
trans-N-feruloyltyramine and cis-N-feruloyl- and myricetin-3-O-glucopyranoside (Lin et al.
tyramine); and 11 flavonoids (quercetin, kaemp- 2009a, b). From the ethyl acetate-n-butanol-water
ferol, luteolin, quercetin 3-O-glucopyranoside, (4:1:5, v/v) extract of lotus leaves, 5 mg of
kaempferol 3-O-glucopyranoside, chrysoeriol quercetin-3-O-sambubioside [quercetin-3-O-β-d-
7-O-glucopyranoside, taxifolin, epitaxifolin, xylopyranosyl (1 → 2)-β-d-glucopyranoside}
5,7,3′,5′-tetrahydroxyflavanone, (–)-catechin and with a purity of 98.6 % was obtained (Deng et al.
elephantorrhizol) (Ahn et al. 2013). 2009a). From lotus leaves, one new compound,
The following compounds quercetin 3-O-α- 24(R)-ethylcholest-6-ene-5α-ol-3-O-β-d-gluco-
arabinopyranosyl-(1 → 2)-β-galactopyranoside, pyranoside, along with 11 known compounds
rutin, (+)-catechin, hyperoside, isoquercitrin, including aporphine alkaloids, diterpenes, a ste-
quercetin and astragalin were isolated from water roid glycoside and a flavonoid glycoside were iso-
lotus leaf ethanol extract (Ohkoshi et al. 2007). lated (Agnihotri et al. 2008). 10-Eicosanol and
Three flavonoid compounds hyperin, isoquercetin 7,11,15-trimethyl-2-hexadecanone were reported
and astragalin were obtained from lotus leaves for the first time. Six compounds were detected
with over 97 % recovery (Tian et al. 2007). In the and characterised in lotus leaf ethanol extract,
ethyl acetate lotus leaf fraction, quercetin 3-O-β- one as catechin glycoside and five as flavonoid
d-glucopyranoside and quercetin 3-O-β-d- glycoside derivatives: miricitrin-3-O-glucoside,
glucuronopyranoside were found as its two major hyperin, isoquercitrin, quercetin-3-O-rhamnoside
components, as well as quercetin 3-O-β-d- and astragalin (Huang et al. 2010b).
galactopyranoside as a minor compound (Jung Six compounds were detected and character-
et al. 2008). A total of 4.6 mg of isoquercitrin, ised in lotus leaf ethanol extract, one as catechin
9.1 mg of hyperoside and 3.0 mg of astragalin glycoside and five as flavonoid glycoside deriva-
with the purity of 95.8 %, 97.5 % and 98.3 %, tives: miricitrin-3-O-glucoside, hyperin, isoquer-
respectively, were obtained from lotus leaf extract citrin, quercetin-3-O-rhamnoside and astragalin
(Deng et al. 2009b). Six flavonoids, quercetin, (Huang et al. 2010b).
rutin, quercetin 3-O-β-d-galactopyranoside From 150 mg of the crude lotus leaf sample,
(Qc-3-Gal), quercetin 3-O-β-d-glucopyranoside 6.1 mg of quercetin-3-O-β-d-glucuronide,
(Qc-3-Glc), quercetin 3-O-β-d-glucuronide 14.8 mg of myricetin-3-O-β-d-glucopyranoside
(Qc-3-Gln) and quercetin 3-O-α-arabinopy- and 20. 2 mg of astragalin were obtained with
ranosyl-(1 → 2)-β-galactopyranoside (Qc-3- purities of 97.0 %, 95.4 % and 96.3 %, respec-
AraGal), were isolated from the methanol leaf tively (Xu et al. 2011c).
extract (Goo et al. 2009). Among them, Qc-3-Glc The polyphenols identified in lotus leaves
and Qc-3-Gln were found to be major compo- consisted primarily of gallic acid, rutin and quer-
nents in the methanol leaf extract. Five cetin (Yang et al. 2011). Quercetin-3-O-β-d-
norsesquiterpenes ((E)-3-hydroxymegastigm-7- glucuronide, hyperoside and isoquercitrin were
en-9-one, (3S,5R,6S,7E)-megastigma-7-ene- the most dominant flavonoids in lotus leaves (Do
3,5,6,9-tetrol, dendranthemoside B, icariside B2 et al. 2012). Twenty-five compounds were iso-
and sedumoside F1), four flavonoids (luteolin, lated from the 70 % ethanol lotus leaf extract and
quercetin 3-O-β-d-glucuronide, quercetin 3-O-β- identified as 10-octacosanol, β-sitosterol,
d-glucoside and isorhamnetin 3-O-rutinoside), 1-undecanol, 1-eicosanol, daucosterol,
two triterpenes (alphitolic acid and maslinic 6′-hydroxy-4,4′-dimethoxychalcone, 3,7,8-trim
acid) and one alkaloid (N-methylasimilobine) ethoxy-1-hydroxy-xanthone, rhamnetin-3-O-
were isolated from the methanol lotus leaf β-d-glucopyranoside, chrysoeriol-7-O-β-d-
extract (Kim et al. 2009). Seven flavonoids were glucoside, quercetin-3-O-β-d-glucopyranoside,
isolated from lotus leaves and identified as cate- quercetin-3-O-α-l-rhamnopyranosyl, hypero-
chin, quercetin, quercetin-3-O-glucopyranoside, side, quercetin-3-O-rutinoside, astragalin,
Nelumbo nucifera 73

isorhamnetin-3- O -α- l - rhamnopyranosyl- kaempferol, kaempferol 3-O-β-d-

(1 → 6)-[α-d-lyxopyranosyl-(1 → 2)-β- glucuronopyranosyl methylester, kaempferol
d-glucopyranoside], isorhamnetin-3-O -α- d - 3-O-β-d-glucopyranoside, kaempferol 3-O-β-d-
lyxopyranosyl-(1 → 2)-β- d - glucopyranoside, galactopyranoside, myricetin 3′,5′-dimethylether
isorhamnetin-3- O -β- d - glucopyranoside, 3-O-β-d-glucopyranoside, kaempferol 3-O-α-l-
isorhamnetin-3- O -α- l - rhamnopyranosyl- rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside
(1 → 6)-β-d-glucopyranoside, quercetin, kaemp- and kaempferol 3-O-β-d-glucuronopyranoside,
ferol, dehydronuciferine, roemerine, stigmast-7- along with β-sitosterol glucopyranoside, were
en-3-O-β-d-glucopyranoside, stigmast-7-en-3β-ol isolated from lotus stamens (Jung et al. 2003).
and benzene-1,2-diol (Zhao et al. 2013b. Mature Thirteen flavonoids including kaempferol and
leaf pulp (771.79 mg/100 g FW (fresh weight)) and seven of its glycosides, myricetin 3′,5′-dim-
young leaves (650.67 mg/100 g FW) of lotus had ethylether 3-O-β-d-glucopyranoside, quercetin
higher total flavonoid amount than flower stamens 3-O-β-d-glucopyranoside and two isorhamnetin
and petals, but leaf stalks had lower (<40 mg/100 g glycosides as well as four non-flavonoid com-
FW) (Chen et al. 2012b). Young leaves, mature leaf pounds (adenine, myo-inositol, arbutin and
pulps and mature leaf veins contained similar range β-sitosterol glucopyranoside) were isolated from
of flavonoids in varying concentrations: myricetin the ethyl acetate-soluble fraction of lotus stamens
3-O-glucoside, quercetin 3-O-arabinopyranosyl- (Lim et al. 2006). Polyphenols found in Korean
(1 → 2)-galactopyranoside, quercetin 3-O-rhamno- traditional lotus liquor (Yunyupju) made from
pyranosyl-(1 → 6)-glucopyranoside (rutin), quer- lotus blossom and leaves were identified as cate-
cetin 3-O-galactoside (hyperoside), quercetin chin, rutin, quercitrin, myricetin and quercetin
3-O-glucoside (isoquercitrin), quercetin 3-O- (Lee et al. 2005). β-carotene concentrations in
glucuronide, kaempferol 3-O-galactoside, kaemp- lotus stamens ranged from 6.50 to 58.50 μg/mL
ferol 3-O-glucoside (astragalin), isorhamnetin (Phonkot et al. 2010). The β-carotene of the four
3-O-glucoside, kaempferol 3-O-glucuronide, dios- lotus varieties was 465.77–1150.80 mg%. Fifteen
metin 7-O-hexose and quercetin; leaf stalks had flavonoids comprising anthocyanins and flavo-
similar flavonoids plus diosmetin. Thirteen flavo- nols were identified in lotus petals: delphinidin
noids were simultaneously separated and identified 3-O-glucoside, cyanidin 3-O-glucoside, petuni-
from lotus leaf methanol–water extract (Chen et al. din 3-O-glucoside, peonidin 3-O-glucoside, mal-
2012c). From lotus leaves, 13 megastigmanes, vidin 3-O-glucoside and quercetin 3-O-rutinoside,
including a new megastigmane, nelumnucifoside A quercetin 3-O-galactoside, quercetin 3-O-
and a new eudesmane sesquiterpene, nelumnucifo- glucuronide, kaempferol 3-O-robinobioside,
side B, eight alkaloids and 11 flavonoids were iso- kaempferol 3-O-galactoside, kaempferol 3-O-
lated (Ahn et al. 2013). glucoside, kaempferol 3-O-glucuronide, isorh-
amnetin 3-O-rutinoside, syringetin 3-O-hexose
and kaempferol 3-O-pentose (Yang et al. 2009).
Flower Phytochemicals From 125 mg crude lotus petal sample, 5.0 mg
syringetin-3-O-β-d-glucoside, 6.5 mg quercetin-
The flavonoid kaempferol-3-galactorhamno- 3-O-β-d-glucoside, 12.8 mg isorhamnetin-
glycoside was isolated from lotus flower (Rahman 3-O-β-d-glucoside and 32.5 mg kaempferol-3-O-
et al. 1962). Two new isorhamnetin glycosides, β-d-glucoside were obtained (Guo et al. 2010). A
designated as nelumboroside A and nelumboro- new β-cyclogeraniol diglycoside, nuciferoside,
side B, as well as known isorhamnetin glucoside with the chemical structure 1-hydroxy-
and isorhamnetin rutinoside were isolated from methyl-2,6,6-trimethyl-1-cyclohexene 9-O-β-d-
the n-BuOH fraction of N. nucifera stamens glucopyranosyl-(1 → 6)-β- d - glucopyranoside,
(Hyun et al. 2006). Seven known flavonoids, along with four known components, cycloartenol,
74 Nelumbonaceae

p-hydroxybenzoic acid, vanilloloside and ferol 3-O-robinobioside, quercetin 3-O-glucuronide,

5′-O-methyladenosine, were isolated from the kaempferol 3-O-galactoside, isorhamnetin
n-BuOH fraction of lotus stamens (Jung 3-O-rutinoside, kaempferol 3-O-glucoside (astraga-
et al. 2010b). lin), syringetin 3-O-glucoside, isorhamnetin
A total of five anthocyanins and 14 flavones 3-O-glucoside and kaempferol 3-O-glucuronide.
and flavonols were detected and quantified in108 Flower pistils had almost similar range plus dios-
lotus cultivars with red, pink, yellow, white and metin 7-O-hexose but minus kaempferol
red/white pied petal colours (Deng et al. 2013). In 3-O-robinobioside. Flower stalks had almost simi-
general, the yellow, white and pied species hardly lar range plus 16 kaempferol 7-O-glucoside, 17
contained any anthocyanins; red cultivars con- diosmetin 7-O-hexose and diosmetin but without
tained more than pink cultivars. Among the five kaempferol 3-O-robinobioside and isorhamnetin
anthocyanins, malvidin 3-O-glucoside was the 3-O-rutinoside. A total of five anthocyanins
most abundant one in all the cultivars that contain (anthocyanins delphinidin 3-O-glucoside, cyani-
anthocyanin. The 14 flavones and flavonols din 3-O-glucoside, petunidin 3-O-glucoside,
belonged to four groups based on their aglycones. peonidin 3-O-glucoside and malvidin 3-O-
Except for the yellow cultivars, kaempferol deriva- glucoside) and 20 non-anthocyanin flavonoids (3
tives were the most abundant one. Flavones and myricetin glycosides [myricetin 3-O-galactoside,
flavonols, deemed as accessory pigments, appeared myricetin 3-O-glucoside, myricetin 3-O-glucu-
to contribute little to the yellow colour. Most of the ronide], 6 quercetin glycosides [quercetin 3-O-
cultivars could be sorted into three groups (A, B arabinopyranosyl-(1 → 2)-galactopyranoside,
and C). Group A exclusively contained mostly red quercetin 3-O-rhamnopyranosyl-(1 → 6)-glucopy-
and several pink cultivars due to high content of ranoside, quercetin 3-O-galactoside,quercetin
total anthocyanins. Group B contained most of the 3-O-glucoside, quercetin-3-O-glucuronide, quer-
yellow cultivars, small part of pink and pied colour cetin], 5 kaempferol glycosides [kaempferol
cultivars and two white cultivars due to high con- 3-O-robinobioside, kaempferol 3-O-galactoside,
tent of flavones and flavonols. Group C contained kaempferol 3-O-glucoside, kaempferol 3-O-
most of the white cultivars due to low content of glucuronide, kaempferol 7-O-glucoside], 3 isorh-
total anthocyanins, flavones and flavonols. amnetins [isorhamnetin 3-O-rutinoside, isorham-
Anthocyanins in lotus flower petals were identi- netin 3-O-glucoside, isorhamnetin 3-O-
fied as delphinidin 3-O-glucoside, cyanidin glucuronide], 2 diosmetin [diosmetin 7-O-hexose,
3-O-glucoside, peonidin 3-O-glucoside and mal- diosmetin] and syringetin 3-O-glucoside) were
vidin 3-O-glucoside (Li et al. 2014). identified and quantified in the petals of 12 lotus
Flower tissues including flower petals, sta- genotypes (Chen et al. 2013).
mens, pistils and, especially, reproductive tissue Fatty acid and sterol content of lotus bee pol-
fruit coats had more flavonoid compounds (15– len oil extracted by supercritical CO2 at 30 MPa
17) than leaves (12) (Chen et al. 2012b). Total and 50 °C comprised of the following: fatty acid
flavonoid content in flower stamens was (myristic acid (14:0), 0.8 %; pentadecanoic acid
359.45 mg/100 g FW and 342.97 mg 100/g FW (15:0), 1.32 %; palmitic acid (16:0), 44.78 %;
in flower petals, while leaf flower stalks had palmitoleic acid (16:1), 0.38 %; margaric acid
much less total flavonoid (less than 40 mg/100 g (17:0), 1.71 %; stearic acid (18:0), 2.84 %; oleic
FW). Flower petals and stamens contained simi- acid (18:1), 18.93; linoleic acid (18:2), 14.78 %;
lar range of flavonoids, namely, myricetin α-linolenic acid (18:3), 7.32 %; behenic acid
3-O-galactoside, myricetin 3-O-glucoside, quer- (22:0), 1.43 %; SFA (saturated fatty acids),
cetin 3-O-arabinopyranosyl-(1 → 2)-galactopy- 52.89 %; MUFA (monounsaturated fatty acids),
ranoside, myricetin 3-O-glucuronide, quercetin 19.31; and PUFA (polyunsaturated fatty acids),
3-O-rhamnopyranosyl-(1 → 6)-glucopyranoside 22.19) and sterols (g/kg oil) (campesterol, 1.39 g;
(rutin), quercetin 3-O-galactoside (hyperoside), stigmasterol, 3.97; β-sitosterol, 2.45 g; and
quercetin 3-O-glucoside (isoquercitrin), kaemp- β-amyrin, 2.90 g) (Xu et al. 2011a). The yield of
Nelumbo nucifera 75

oil was 7.18 g/kg; carotenoids, 377.86 mg/kg; galactoside (f13), kaempferol 3-O-robinobioside
squalene, 84.94 mg/kg; and sterols, 490.90 mg/ (f14), isorhamnetin 3-O-rutinoside (f15), kaemp-
kg dry bee pollen. ferol 3-O-β-d-glcuopyranoside (f16), kaempferol
3-O-glucuronide (f17), isorhamnetin 3-O-β-d-
glucopyranoside (f18), isorhamnetin 3-O-glucu-
Aerial Plant Parts Phytochemicals ronide (f19), syringetin 3-O-glucuronide (f20),
apigenin di-C-glucoside (f21), 6-C-glucosyl-8-C-
Alkaloids were found in the leaves, petiole and pentosyl luteolin (f22), 6-C-pentosyl-8-C-glucosyl
cotyledons in the fruit but not in the edible rhi- luteolin (f23), 6-C-glucosyl-8-C-xylosyl api-
zome (Tomita et al. 1961c). All the alkaloids genin (f24), 6-C-xylosyl-8-C-glucosyl api-
found were in a free state. Roemerine and nucif- genin (f25), 6-C-glucosyl-8-C-arabinosyl api-
erine were isolated as the tertiary, non-phenolic genin (schaftoside) (f26), 6-C-arabinosyl-8-C-
bases, and a new base, nornuciferine, as the ter- glucosyl apigenin (isoschaftoside) (f27), quer-
tiary phenolic base, from the leaves. Roemerine cetin 3-O-neohesperidoside (f28), 6-C-gluco-
and nornuciferine were isolated from the petiole syl-8-C-rhamnosyl apigenin (f29), luteolin
but not nuciferine. After methanolic extraction, 7-O-rutinoside (f30), 6-C-rhamnosyl-8-C-glu-
the phenolic content of each part of the plant, cosyl apigenin (f31), isorhamnetin 3-O-neohes-
determined in terms of gallic acid equivalents per peridoside (f32) and diosmetin 7-O-rutinoside
g of dried extract (GAE, mg/g extract), was found (f33). Flower petals and stamens possessed the
to follow the order of leaves (177.7) > de- highest amounts of kaempferol glycoside, spe-
embryonated seeds (92.7) > stamens (83.4) > cially f11 and f17. Additionally f14 and f16 were
embryos (41.0) > rhizomes (21.6). For the flavo- the predominant compounds in petals. The petals
noid contents, determined as mg of (+)-catechin contained f1, f4, f5, f7, f8, f10, f11, f14, f15, f16,
equivalent per g of dried extract (CE, mg/g f17, f18 and f19. The stamens contained f1, f4,
extract), the order was leaves (125.6) > de- f5, f7, f8, f10, f11, f12, f14, f15, f16, f17, f18 and
embryonated seeds (82.9) > stamens (50.3) > f19. The pistil and tori contained f1, f4, f6, f8,
embryos (18.9) > rhizomes (8.5) (Jung et al. f14, f15, f16, f17, f18 and f19. The flower stalks
2008). contained f1, f4, f6, f8, f10, f15, f18 and f19; the
Total flavonoid content was highest in the leaves contained f1, f4, f6, f11, f15, f16, f18 and
leaves 2.06 E3mg/100 g FW, followed by flower f20. Lotus seed pods contained f4, f6, f10, f11,
petals, stamens, plumule 402.9–496 mg/100 g f14, f15, f16, f17, f18 and f19; seed coats con-
FW and seed kernel the lowest 0.67 mg/100 g tained f2, f3, f4, f7, f9, f11, f12, f13, f14, f15, f16,
FW (Li et al. 2014). Thirty-three flavonoids were f17, f18 and f19. The seed kernels contained f9,
found in lotus flowers, seeds and leaves: f11, f12, f15, f16 and f18. Thirteen C-glycosyl
myricetin 3-O-galactoside (f1), luteolin 8-C-β-d- flavones (newly isolated C-glycosides, f7, f9,
glucopyranoside (orientin) (f2), luteolin 6-C-β-d- f21, f22, f23, f24, f25, f26, f27, f29 and f31) and
glucopyranoside (isoorientin) (f3), myricetin six O-glycosides (newly isolated O-glycosides,
3-O-glucuronide (f4), myricetin 3-O-glucoside f20, f28, f30, f32 and f33) within plumules were
(f5), quercetin 3-O-arabinopyranosyl-(1 → 2)- detected.
galactopyranoside (f6), apigenin 8-C-β-d- The extracts of lotus rhizomes, seeds, flowers
glucopyranoside (vitexin) (f7), quercetin 3-O-α- and leaves had been reported to have varied ther-
l-rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside apeutic potential (Mukherjee et al. 2009). Several
(rutin) (f8), apigenin 6-C-β-d-glucopyranoside bioactive compounds had been derived from
(isovitexin) (f9), quercetin 3-O-β-d-galactopy- these plant parts belonging to different chemical
ranoside (hyperoside) (f10), quercetin 3-O- groups, including alkaloids, flavonoids, glyco-
glucuronide (f11), quercetin 3-O-β-d-glucopy- sides, triterpenoid, vitamins, etc., which all
ranoside (isoquercitrin) (f12), kaempferol 3-O- imparting their own therapeutic impact.
76 Nelumbonaceae

Pharmacological studies have demonstrated that values in both DPPH (1,1-diphenyl-2-picryl

lotus herb exhibits various pharmacological hydrazyl) (6.12 μg/ml) and nitric oxide (84.86 μg/
effects, such as anti-hyperlipidaemia, antiobesity, ml) methods; the values were found to be lower
antioxidant, anti-HIV, antimicrobial and anti- than those of rutin, the standard used (Rai et al.
hypoglycaemic activities. 2006). Total phenolic content in the extract was
found to be 7.61 % (w/w). The Fr2 fraction from
lotus seed epicarp exhibited high antioxidant
Antioxidant Activity activity as assessed by 2,2-diphenyl-1-
picrylhydrazyl (DPPH) radical scavenging, the
Lotus Seed/Seedpod/Seedlings b-carotene bleaching method and hydroxyl radi-
Lotus seed water, ethyl acetate and hexane cal and hydrogen peroxide scavenging ability
extracts could inhibit nitric oxide accumulation using the chemiluminescence method (Kredy
in LPS-activated RAW 264.7 cells (Yen et al. et al. 2010). This antioxidant potential in terms of
2006). The extracts in the range of 0.01–0.2 mg/ IC50 values was 5.48, 40 and 0.62 g (dry Fr2)/mL
ml showed a dose-dependent inhibitory effect on on DPPH radicals, hydroxyl radicals and hydro-
the accumulation of nitric oxide upon decompo- gen peroxide, respectively. The Fr2 also exhib-
sition of sodium nitroprusside (SNP). The ited antioxidant property in the b-carotene
potency of inhibitory activity was in the order bleaching assay. In total, it possessed high levels
ethyl acetate > water > hexane. The results of the of flavonol compounds with high antioxidant
comet assay indicated that the three extracts potential. The DPPH free radical scavenging
could inhibit DNA damage in macrophage RAW activities of lotus seed extract (LSE) increased in
264.7 cells induced by SNP. Further, the three a concentration-dependent manner (Sung et al.
extracts, at 0.2 mg/ml, showed 63 %, 59 % and 2011). Mouse embryonic fibroblast cells, dam-
38 % inhibition of DNA damage in macrophage aged by oxidative stress, decreased their viability
RAW 264.7 cells induced by peroxynitrite, following increasing concentration of H2O2, but
respectively. All extracts tested were found to be the cotreatment of ethyl acetate fraction of
potent peroxynitrite scavengers, capable of pre- lotus seed extract and H2O2 resulted in an
venting the nitration of tyrosine. The data increase in cell growth, by about 25 %, compared
obtained suggested that lotus seed extracts might to the cells treated with H2O2. The ethyl acetate
act as chemopreventers through reduction of fraction of LSE inhibited the cytotoxicity
excess amounts of nitric oxide. Lotus seeds pos- induced by H2O2 in a concentration-dependent
sessed higher temperature tolerances than maize manner. The results suggested that LSE
seeds (Ding et al. 2008). Germination percentage inhibited the cytotoxicity induced by H2O2 and
of maize (Zea mays ‘Huangbaogu’) seeds was exerted a protective effect on mouse embryonic
zero after they were treated at 100 oC for 15 min fibroblast cell against oxidative stress. Hexane
and that of lotus seeds was 13.5 % following the extract of lotus seed oil showed 66 % inhibition
treatment at 100 oC for 24 h. For lotus seeds, (1) of DPPH radical with IC50 of 15 μgml
activities of superoxide dismutase (SOD) and (Anitha and Arunkumar 2012). It also exhibited
glutathione reductase (GR) of axes and cotyle- the highest ABTS assay activity with IC50 of
dons and of catalase (CAT) of axes increased dur- 124 μg/ml.
ing the early phase of treatment at 100 oC and Studies showed that 0.1 % procyanidins from
then decreased and (2) activities of ascorbate per- lotus seed pod had a strong antioxidant activity in
oxidase (APX) and dehydroascorbate reductase a soybean oil system, better than BHT at the
(DHAR) of axes and cotyledons and of CAT of same concentration; inhibited lipoxygenase
cotyledons gradually decreased with increasing activity by >90 % at a concentration of 62.5 mug/
treatment time at 100 oC. The hydroalcoholic mL, with an IC50 value of 21.6 mug/mL; and had
lotus seed extract exhibited strong free radical IC50 inhibitory value rate to *OH of 10.5 mg/L
scavenging activity as evidenced by the low IC50 and a scavenging effect on O2* of 17.6 mg/L
Nelumbo nucifera 77

(Ling et al. 2005). DPPH and ABTS scavenging Lotus Leaf

capacities of the ethyl acetate fraction of lotus A dose-dependent protective effect against reac-
seed (LS) were 94.6 and 91.9 % and those of the tive oxygen species (ROS)-induced cytotoxicity
water fraction of lotus seedpod (LSP) were 94.5 was observed when Caco-2 cells were treated
and 95.2 % at 0.8 mg/mL (Kim and Shin 2012). with 10 mM H2O2 in combination with the meth-
The ethyl acetate fraction of LS and the water anol extract of the lotus leaf (0.1–0.3 mg/ml) (Wu
fraction of LSP also showed high ferric reducing et al. 2003). However, no significant effect was
ability of plasma (FRAP). The high antioxidant found when cotreating Caco-2 cells with 10 mM
capacities of LS and LSP may be due to their fla- H2O2 and α-tocopherol. In-vitro assay revealed
vonoid and proanthocyanidin contents. The that lotus leaf extract exhibited scavenging activi-
n-butanol fraction of lotus receptacle showed the ties on free radicals and hydroxyl radicals and
highest total phenolic content (607.6 mg/g gallic metal binding ability as well as reducing power,
acid equivalents), total flavonoid content which explained partly the mechanism behind
(862.7 mg/g rutin equivalents) and total proan- the extract’s ability to protect cells from oxida-
thocyanidin content (331.0 mg/g catechin equiv- tive damage. Moreover, the extract also exhibited
alents), accompanied with the highest antioxidant concentration-dependent antioxidant activities
activity compared to other fractions in the against haemoglobin-induced linoleic acid per-
1,1-diphenyl-2-picrylhydrazyl (DPPH) and 2,2′- oxidation and Fenton reaction-mediated plasmid
azino-bis-(3-ethylbenzthiazoline-6- sulphonic DNA oxidation. Lotus leaf methanol extract
acid) (ABTS) radical scavenging assays (Wu exhibited a pronounced activity in the
et al. 2012). Among the five isolated flavone gly- 1,1-diphenyl-2-picrylhydrazyl (DPPH) scaveng-
cosides, hyperoside, isoquercitrin and quercetin- ing assay with an IC50 of 90 μg/ml, compared
3-O-β-d-glucuronide demonstrated significant with an IC50 of 30 μg/ml for the butylated
DPPH and ABTS radical scavenging activity, hydroxytoluene (BHT) control (Hutadilok-
with IC50 values of 8.9, 5.2 and 7.5 for DPPH Towatana et al. 2006). The same extract was also
and 114.2, 112.8 and 172.5 μg/mL for ABTS, found to be the most potent in removing the
respectively. superoxide anion (O 2) radical and in inhibiting
The activity of superoxide dismutase (SOD), the 2,2′-azo-bis-(2-amidinopropane) dihydrochlo-
dehydroascorbate reductase (DHAR) and gluta- ride (AAPH)-induced erythrocyte haemolysis
thione reductase (GR) was lower in lotus seed- and lipid peroxidation in a rat brain
lings germinated under water (submerged homogenate.
hypoxic condition) in darkness (SD seedlings) The ethyl acetate and n-butanol fractions of
than those found in seedlings germinated in air lotus leaf methanol extract exhibited greater
and darkness (AD seedlings) (Ushimaru et al. capacity to scavenge DPPH radical, delayed low-
2001). In contrast, ascorbate peroxidase activity density lipoprotein (LDL) oxidation and had
was higher in SD seedlings, and the activity of higher antioxidative contents than the water frac-
catalase and monodehydroascorbate reductase tion (Lin et al. 2009a). Among its constituents,
(MDAR) in SD seedlings was nearly the same as quercetin and its glycosides quercetin-3-O-
in AD seedlings. When SD seedlings were glucopyranoside, quercetin-3-O-glucuronide and
exposed to air, the activity of SOD, DHAR and quercetin-3-O-galactopyranoside exerted potent
GR increased, while the activity of catalase and inhibition of LDL oxidation, whereas myricetin-
MDAR decreased. Seven electrophoretically dis- 3-O-glucopyranoside (7) showed stronger DPPH
tinct SOD isozymes were detectable in N. scavenging activity. Jeong et al. (2009) reported
nucifera. The levels of plastidic Cu, Zn-SODs that radical scavenging activities and total poly-
and Fe-SOD in SD seedlings were comparable phenols of the γ-irradiated lotus leaf ethanol
with those found in AD seedlings, which may extract were not observed to be significantly dif-
reflect the maintenance of green plastids in SD ferent. However, γ-irradiation significantly
seedlings as well as in AD seedlings. increased the Hunter colour L*-value at doses of
78 Nelumbonaceae

20 and 50 kGy, while the Hunter colour b*-val- petroleum extracts had comparable AAC with
ues were decreased under the same conditions. BHA by the beta-carotene bleaching assay.
Lotus leaf extract exhibited 2-diphenyl-1- Lotus rhizome knot extract exhibited high
picrylhydrazyl hydrate (DPPH) free radical antioxidative capacity, as measured by1-
scavenging activities increased in a concentra- diphenyl-2-picrylhydrazyl and 2,2′-azino
tion-dependent manner (Ha et al. 2010). Mouse (3-ethylbenzothioazolino-6-sulfonate) and mea-
embryonic fibroblast (MEF) cells, damaged by sured by electron spin resonance (ESR) trapping
oxidative stress, decreased their viability follow- of the transient carbon-centred 1-hydroxyethyl
ing increasing concentration of H2O2, but the radical (generated in a Fenton-type reaction) (Hu
cotreatment of n-butanol fraction of the leaf and Skibsted 2007). Whole rhizome, however,
extract and H2O2 resulted in an increase in cell only showed a significantly high scavenging
growth, by about 25 %, compared to the cells activity for small carbon-centred radicals, as
treated with H2O2. At 100 μg/ml extract concen- measured by the ESR method. Total phenol con-
tration, lotus leaf, flower and seed extracts tent in the plant extract correlated with the anti-
showed 78 %, 78 % and 80 % DPPH free radical oxidant capacity, except for the scavenging of
scavenging activity (Kim et al. 2011). The IC50 carbon-centred radical. Studies found that
values (μg/ml) of the antioxidation effects were gamma radiation treatment significantly reduced
flower, 5 μg; seed, 5 μg; leaf, 29 μg; root, 150 μg; microbial load, enhanced the DPPH scavenging
stem(rhizome), 162 μg; and vitamin C, 5 μg. activity and increased the storability of the irradi-
The amounts of phenolics, flavonoids and pro- ated lotus rhizome samples (Khattak et al. 2009).
anthocyanidins in the lotus leaf extracts from The treated samples were also acceptable sen-
various areas in China varied widely, ranging sorically. The extraction yield and phenolic con-
from 354 to 487 mg/g gallic acid equivalents, tents increased with the increase of radiation
from 172 to 236 mg/g rutin equivalents and from dose.
124 to 179 mg/g catechin equivalents, respec- The lotus rhizome showed the strongest anti-
tively (Huang et al. 2010a). All of the extracts oxidant activity in both assays of lipid peroxida-
had strong antioxidant activity in comparison to tion and inhibition of lysis of erythrocytes
the standard compounds butylated hydroxytolu- induced by peroxyl radicals (Jiang et al. 2010).
ene and vitamin C. Wild lotus samples from Of three fractions, L2 fraction showed the high-
Baiyangdian Lake and Weishan Lake exhibited a est antioxidant activity and was further fraction-
stronger free radical scavenging effect and greater ated, and L2c showed the strongest activity in
reducing power than the cultivated samples, but inhibiting haemolysis of erythrocytes, and on
no such differences were observed in the inhibi- further purification, L2c-3 was purified and
tion of lipid oxidation. identified as tryptophan. Its inhibitory concen-
tration of 50 % (IC50) value in inhibiting hae-
Lotus Rhizome molysis of erythrocytes was 156.3 μg/ml (i.e.
Methanol extract of lotus rhizome powder 765.4 μM).
showed the highest extract yield among all of sol-
vents (Yang et al. 2007). Although acetone Lotus Flower
extract had the highest total phenolics content, Ethanol lotus flower extract exhibited higher
methanol extract had the highest total phenolics antioxidant activity in-vitro (lower IC50 value)
recovery from lotus powder (20.1 mg catechin than vitamin C (Gayathri et al. 2009). A metha-
equivalents/100 g lotus powder). Extract of either nol extract of lotus stamens showed strong anti-
methanol or acetone demonstrated the highest oxidant activity in the peroxynitrite (ONOO-)
DPPH scavenging activity at both 66.7 and system and marginal activity in the DPPH and
133.3 mg/L. All extracts exhibited higher anti- total reactive oxygen species (ROS) systems
oxidant activity coefficient (AAC) than that of (Jung et al. 2003). After fractionation, the ethyl
ascorbic acid; furthermore, dichloromethane and acetate fraction exhibited strong antioxidant
Nelumbo nucifera 79

activity in all the model systems tested, and Anticancer Activity

among its constituents, kaempferol exhibited
good antioxidant activities in all the model sys- Oxoushinsunine, a cytotoxic alkaloid isolated
tems tested. Compounds kaempferol 3-O-β-d- from lotus receptacle, showed tumour inhibitory
glucuronopyranosyl methylester and kaempferol activity against nasopharynx carcinoma (Yang
3-O-β-d-glucuronopyranoside showed scaveng- et al. 1972). Luteolin, alphitolic acid, maslinic
ing activities in the DPPH and ONOO- tests, acid and N-methylasimilobine, isolated from
while compounds kaempferol 3-O-β-d-glucopy- lotus leaves, exhibited considerable cytotoxicity
ranoside and kaempferol 3-O-β-d-galactopy- against four human cancer A549 (non-small cell
ranoside were only active in the ONOO-test. lung adenocarcinoma), SK-OV-3 (ovarian can-
Conversely, compound β-sitosterol glucopyrano- cer), SK-MEL-2 (skin melanoma) and HCT15
side showed no activities in any of the model sys- (colon cancer) cell lines in-vitro using a SRB
tems tested. (sulforhodamine B) bioassay (Kim et al. 2009).
All isolated isorhamnetin glycosides, nelum- Among Korean white lotus cultivar rhizomes,
boroside A, nelumboroside B, isorhamnetin glu- the highest contents of polyphenols (7.95 mg of
coside and isorhamnetin rutinoside, from lotus GAE/g DW) and the highest levels of antioxi-
stamens showed marked antioxidant activities in dant [by 2,2-azino-bis(3-ethylbenzthiazoline-6-
the DPPH and peroxynitrite (ONOO)– assays sulfonic acid) and 1,1-diphenyl-2-picrylhydrazyl
(Hyun et al. 2006). The DPPH antioxidant activi- assays, 54.27 and 21.98 μM Trolox equivalents/g
ties (IC50) of methanol stamen solution of four DW, respectively] and antiproliferative activities
lotus varieties were 68.30 6.30, 62.22 4.00, 31.60 on both human cancer cell lines (Calu-6 for
3.40 and 40.90 1.50 g/mL, respectively, whereas human pulmonary carcinoma and SMU-601 for
those of the mixed-solvent solution were 2.21 human gastric carcinoma, 59.75 % and 71.21 %
0.06, 2.23 0.05, 1.29 0.02 and 1.83 0.07 mg/mL cell viability, respectively) were found in the
(Phonkot et al. 2008). Chungyang cultivar (Park et al. 2009). Neferine
possessed a potent growth-inhibitory effect on
Lotus Liquor human osteosarcoma cells, but not on non-
The antioxidant activities of lotus liquor neoplastic human osteoblast cells (Zhang et al.
(Yunyupju) made from lotus blossom and leaves 2012). The inhibitory effect of neferine on
were dose dependent and reached a plateau human osteosarcoma cells was largely attributed
(about 80 % inhibition) when the concentration to cell cycle arrest at G1. The induction of G1
of lotus liquor exceeded 25 μg in a modified lin- arrest was p21(WAF1/CIP1) dependent, but was
oleic acid peroxidation induced by haemoglobin independent of p53 or RB (retinoblastoma-asso-
(Lee et al. 2005). The concentrations to attain ciated protein). Neferine treatment led to an
one absorbance unit at 700 nm were 23.6 μg for increased phosphorylation of p21 at Ser130 that
butylated hydroxytoluene (BHT) and 45.7 μg was dependent on MAPK and JNK. Compounds
for lotus liquor. The scavenging activities of liriodenine, nuciferine, nornuciferine, armepa-
DPPH exerted by lotus liquor and α-tocopherol vine, N-norarmepavine and N-methylcoclaurine,
were obtained and the IC50 values were esti- from lotus rhizome, showed significant cytotoxic
mated at 17.9 μg for lotus liquor and 5.6 μg for activities against HL-60 carcinoma cell line with
α-tocopherol. The maximum scavenging activ- inhibitory ratios of 51.36 %, 59.09 %, 52.51 %,
ity on hydroxyl radicals (40 %) could be 53.93 %, 51.43 % and 64.31 % at concentration
achieved when lotus liquor was more than of 1 × 10−5 mol/L, respectively (Duan et al.
500 μg. Lotus liquor also exhibited potent 2013). Neferine, from lotus embryo, induced
superoxide radical scavenging activity, with autophagy in A549 human lung adenocarcinoma
value of 0.93 unit/mg as superoxide dismutase epithelial cells by inhibiting PI3K/Akt/mTOR
equivalents. The IC50 was estimated at 1.07 mg pathway and reactive oxygen species hypergen-
for lotus liquor. eration (Poornima et al. 2013b). Neferine
80 Nelumbonaceae

inhibited human lung cancer A549 cells by Antiviral Activity

inducing apoptosis in a dose-dependent manner
with the hypergeneration of reactive oxygen spe- The hot water extract of N. nucifera at a concen-
cies, activation of MAPKs, lipid peroxidation, tration of 125 μg/ml inhibited the activity of
depletion of cellular antioxidant pool, loss of Moloney murine leukaemia virus reverse tran-
mitochondrial membrane potential and intracel- scriptase in-vitro with a relative inhibitory ratio
lular calcium accumulation (Poornima et al. (IR) over 50 % (Suthienkul et al. 1993). The
2014). Furthermore, neferine treatment caused methanol extract exhibited an IR value of 54 %.
the inhibition of nuclear factor kappaB and The aporphine alkaloid, (–)-nuciferine HCl, was
Bcl2, upregulation of Bax and Bad, release of found to be active in-vitro against human poliovi-
cytochrome C, activation of caspase cascade rus (Boustie et al. 1998). (+)-1(R)-Coclaurine (1)
and DNA fragmentation. In addition, neferine and (−)-1(S)-norcoclaurine (3), together with
could induce p53 and its effector protein p21 quercetin 3-O-β-d-glucuronide (4), were isolated
and downregulation of cell cycle regulatory pro- from lotus leaves and identified as anti-HIV prin-
tein cyclin D1 thereby inducing G1 cell cycle ciples (Kashiwada et al. 2005). Compounds 1 and
arrest. Neferine was also found to exert cytotox- 3 demonstrated potent anti-HIV activity with
icity on liver cancer cells HepG2 in a dose- EC50 values of 0.8 and <0.8 μg/mL, respectively,
dependent manner (Poornima et al. 2013a). The and therapeutic index (TI) values of >125 and
data suggested that mitochondrial-mediated >25, respectively. Compound 4 was less potent
ROS (reactive oxygen species) generation (EC50 2 μg/mL). Also other liensinine (14), nefer-
induced by neferine led to caspase-dependent ine (15) and isoliensinine, isolated from lotus
apoptosis in HepG2 cells. In another study, leaves, also showed potent anti-HIV activities
neferine exhibited cytotoxicity against hepato- with EC50 values of <0.8 μg/mL and TI values of
cellular carcinoma Hep3B cells (Yoon et al. >9.9, >8.6 and >6.5, respectively. Nuciferine
2013). Neferine induced cell cycle arrest, (12), an aporphine alkaloid, had an EC50 value
endoplasmic reticulum (ER) stress and apopto- of 0.8 μg/mL and TI of 36. Ethanol lotus seed
sis, acting through multiple signalling cascades. extract significantly blocked Herpes simplex
The significant reduction of the migration in virus (HSV-1) multiplication in HeLa cells with-
Hep3B cells and the capillary tube-like out apparent cytotoxicity (Kuo et al. 2005). The
formation of HUVECs by neferine were also mechanisms of antiviral action of the extract
determined. appeared to be mediated, at least in part, through
Oral administration of hydroethanolic extracts inhibition of immediate early transcripts, such as
of pink and white lotus flowers exerted antitumor ICP0 and ICP4 mRNA and then blocking of all
effect by modulating lipid peroxidation and aug- downstream viral products accumulation and
menting antioxidant defence systems in Ehrlich progeny HSV-1 production.
ascites carcinoma bearing mice (Brindha and The antioxidant components L2f-2, L2f-3
Arthi 2010b). Lotus white flower possessed (micromolecular) and LB2 (macromolecular)
potent antioxidant and antitumor activity when from lotus rhizomes strongly inhibited HIV-1 RT
compared with lotus pink flowers. Flavonoid- (reverse transcriptase) and integrase (Jiang et al.
enriched lotus leaf extract exhibited antiprolifer- 2011). L2f-2 was identified as (±)- gallocatechin,
ative effect on MCF-7 human breast cancer cell L2f-3 was identified as (−) catechin and LB2 was
line in-vitro and in-vivo (Yang et al. 2011). a polysaccharide-protein complex. LB2 inhibited
MCF-7 cells treated with the extract were arrested HIV-1 RT HIV-1 RT with an IC50 value of
at the G0/G1 phase (approximately 86.54 %). In 33.7 μM. It also exhibited the highest HIV-1
xenograft nude mice inoculated with MCF-7 3′-processing inhibitory activity with an IC50
cells, treatment with the extract effectively value of 5.28 μM. Both L2f-2 and L2f-3 upregu-
reduced tumour volume and tumour weight com- lated the expression of IL-2 (interleukin-2) and
pared to control. downregulated IL-10, while LB2 exhibited posi-
Nelumbo nucifera 81

tive regulation on IL-2, IL-4 and IL-10. 3-O-glucuronide (Q3GA), a major quercetin
Furthermore, L2f-3 and LB2 might inhibit HIV-1 metabolite after absorption from the digestive
directly by downregulating TNFα (tumour necro- tract, enhanced ATP-binding cassette, subfam-
sis factor α). ily A and member 1 (ABCA1) expression, in-
vitro, via LXRα in macrophages. ABCA1 is a
crucial cholesterol transporter involved in
Antiobesity/Antihyperlipidaemic/ reverse cholesterol transport to produce high-
Anticholesterolemic Activities density lipoprotein-cholesterol.

In-Vitro Studies Animal Studies

Nuciferine, from lotus leaves, dose-dependently Oral administration of aqueous extracts of lotus
inhibited the synthesis of cholesterol and choles- leaves (Kayo), leaves and stem of Lonicera
terol esterase activity in Bel-7402 cells and aug- japonica (nindo) and stems of Akebia quinata
mented low-density lipoprotein receptor (mokutsu) to rats loaded with a high-fat diet con-
expression to reduce blood lipid levels (Han et al. taining 1.5 % cholesterol and 1.0 % cholic acid
2008). significantly decreased serum total cholesterol
Studies showed that incubation of preadipo- (TC), free cholesterol (FC) and phospholipid
cytes with lotus leaf extract solution signifi- (Onishi et al. 1984). The overall effects in sup-
cantly decreased triglyceride accumulation pressing serum lipid elevation were in the order
during adipogenesis without affecting cell via- of Nindo > Mokutsu > Kayo. A decoction of
bility (Siegner et al. 2010). Compared to con- Nelumbo nucifera was tested and found to cause
trols, adipocytes incubated with lotus leaf a reduction of triglyceride and cholesterol
extract solution exhibited a significant increase (LaCour et al. 1995).
in lipolysis activity. It was also demonstrated Studies showed that hepatic and serum total
that a combination of lotus leaf extract and cholesterol levels were significantly lowered on
l-carnitine reduced triglyceride accumulation in day 40 in F1B hamster given lotus leaf extract
human (pre)adipocytes by affecting different extracts (200, 400 mg/kg/day) when compared
processes during the adipocyte life cycle. This with a control value of high-fat atherogenic
combination might represent a treatment option diet-fed hamsters (Kim et al. 2005). But the HDL
for obesity-related diseases. Both methanol and levels were not changed between control and
successive water extracts of lotus petals had an treated lotus leaf groups.
effect on inhibition of lipid storage in adipo- Treatment of high-fat diet-induced obese mice
cytes and on increasing lipolysis (Velusami with N. nucifera leaf extract for 5 weeks caused a
et al. 2013). Lotus petal methanol extract exhib- concentration-dependent inhibition of the activi-
ited concentration-dependent inhibitory effect ties of α-amylase and lipase and upregulated lipid
on lipase activity with an IC50 value of 47 μg/ metabolism and expression of UCP3 mRNA in
mL. Lotus petal extracts showed marked agonist C2C12 myotubes (Ono et al. 2006). The extract
and antagonist activity towards serotonin (5-) prevented the increase in body weight, parame-
and cannabinoid (CNR2) receptors, respec- trial adipose tissue weight and liver triacylglyc-
tively, while it showed no effect towards mela- erol levels in the obese mice. UCP3 mRNA
nocyte concentrating hormone (MCHR1) and expression in skeletal muscle tended to be higher,
melanocortin (MC4R) receptors. Overall, the when mice were administrated by the extract and
methanol extract showed better activity than made to exercise. The results suggested that N.
successive water extract. Lotus leaf extracts nelumbo leaf extract impaired digestion, inhib-
which contained abundant amounts of quercetin ited absorption of lipids and carbohydrates,
glycosides significantly elevated plasma high- accelerated lipid metabolism and upregulated
density lipoprotein-cholesterol (HDLC) in mice energy expenditure and was beneficial for the
(Ohara et al. 2013). They found that quercetin- suppression of obesity.
82 Nelumbonaceae

A 50 % ethanol leaf extract of N. nucifera acetyl-CoA carboxylase and HMGCoA reduc-

stimulated lipolysis in the white adipose tissue tase and increased the phosphorylation of AMP-
(WAT) of mice via the β-adrenergic receptor activated protein kinase in the liver. The results
(β-AR) pathway (Ohkoshi et al. 2007). Dietary suggested that lotus leaf extract targeted lipid-
supplementation of the extract resulted in a sig- regulated enzymes and may be effective in atten-
nificant suppression of body weight gain in A/J uating body lipid accumulation and preventing
mice fed a high-fat diet. Of the compounds iso- obesity. Studies by Du et al. (2010) found that
lated from the leaf extract, quercetin 3-O-α- oral administration of hot water lotus leaf extract
arabinopyranosyl-(1 → 2)-β-galactopyranoside, with taurine supplementation showed antiobesity
(+)-catechin, hyperoside, isoquercitrin and and hypolipidaemic effects in high-fat diet-
astragalin exhibited lipolytic activity, especially induced obese male Sprague–Dawley rats, which
in visceral adipose tissue. The flavonoid-enriched was more effective than lotus leaf hot water
N. nucifera leaf extract supplement effectively extract alone. The combined supplementation
ameliorated the high-fat diet-induced lipid meta- elicited better blood lipid profiles compared to
bolic disorders in hamster such as the significant lotus leaf hot water extract alone. Lotus leaf alka-
increases of body weight, plasma lipid deriva- loid inhibited 3T3-L1 preadipocyte differentia-
tives (triglyceride, total cholesterol and lipopro- tion and ameliorated high-fat diet (HFD)-induced
teins), lipid peroxidation and liver damage obesity and body fat accumulation in rats (Xie
markers (plasma aspartate aminotransferase and et al. 2011). Furthermore, it reduced the body
alanine aminotransferase), as did silymarin and weight, Lee’s index, adipose tissue weight and
simvastatin (Lin et al. 2009b). Moreover, the fla- plasma lipid levels in HFD-induced obese rats.
vonoid-enriched supplement alleviated the high- Oral administration of lotus leaf extract to rats
fat diet-induced accumulation of lipids in liver, fed a high-fat diet significantly decreased body
the findings showing distinguishing mechanisms weight, total cholesterol and triglyceride contents
from the effects of silymarin and simvastatin. in liver tissues and serum levels of total lipids,
Lotus leaf water extract significantly inhibited total cholesterol and triglycerides compared to
hyperlipidaemia and atherosclerosis in rabbits untreated rats fed a high-fat diet (Lee and Lee
fed with high-cholesterol diet (Lee et al. 2010). 2011). Liu et al. (2013) demonstrated that lotus
In rabbits fed with high-cholesterol diet (HCD) leaf flavonoids (NLF) could effectively amelio-
plus 0.5 or 1.0 % of lotus leaf extract, the inhibi- rate hyperlipidaemia and inhibit the key enzymes
tory level of triglyceride was 30 and 46.6 %, and related to type 2 diabetes mellitus in high-fat
low-density lipoprotein-cholesterol (LDL-C) diet-induced hyperlipidaemic rats. In-vitro, NLF
was 45.4 and 45.9 % compared with the HCD- showed high inhibitory activity against porcine
fed rabbits. The extract significantly reduced pancreatic lipase, α-amylase and α-glucosidase
severe atherosclerosis, foam cell formation and with IC50 values of 0.38, 2.20 and 1.86 mg/mL,
smooth muscle cell proliferation and migration in respectively.
the aorta. In the thiobarbituric acid-reacting sub- In-vitro treatment with lotus ethanol seed
stance assay, the concentration of the extract extract resulted in inhibition of lipid accumula-
higher than 0.05 mg/mL, the inhibitory percent- tion and decreased expression of peroxisome
age was about 80 % and inhibited oxidative LDL proliferator-activated receptor gamma (PPARγ),
up to 60 %. glucose transporter 4 (GLUT4) and leptin in cul-
Administration of a flavonoid-enriched lotus tured human adipocytes (You et al. 2014a). In
leaf extract to C57BL/6 mice fed with a high-fat addition, the extract had a beneficial effect,
diet, reduced body weight, body lipid accumula- reducing adipose tissue weights, ameliorating
tion and activities of fatty acid synthase (FAS), blood lipid profile and modulating serum leptin
glutamic oxaloacetic transaminase and glutamic level in rats fed a high-fat diet. Administration of
pyruvic transaminase (Wu et al. 2010). Lotus ethanol lotus rhizome extract resulted in a signifi-
extract also suppressed the expression of FAS, cant decrease in relative weights of adipose tis-
Nelumbo nucifera 83

sues in rats fed a high-fat diet (You et al. 2014b). that lotus leaf extract had potential as an antiobe-
Consumption of a high-fat diet resulted in an sity agent by inhibiting pancreatic lipase and adi-
increase in serum total cholesterol (TC) and tri- pocyte differentiation (Ahn et al. 2013). Among
glyceride (TG) levels; however, administration of its constituents, the alkaloids trans-N-
lotus extract resulted in a decrease in the levels of coumaroyltyramine and cis-N-feruloyltyramine
TC and TG. Administration of the extract also significantly inhibited pancreatic lipase, whereas
resulted in a decrease in the level of serum leptin alkaloids (6R, 6aR)-roemerine-Nβ-oxide and liri-
and insulin and in hepatic thiobarbituric acid odenine showed strong inhibitory effect on adi-
reactive substance content, elevated by a high-fat pocyte differentiation.
diet and caused an increase in superoxide dis-
mutase activity and hepatic glutathione content.
The results suggested that lotus root exerts anti- Antidiabetic Activity
oxidant and antiobesity effects and could be used
as a functional and nutraceutical ingredient in Lotus rhizome extract exhibited hypoglycaemic
combating obesity-related diseases. effect in streptozotocin-induced diabetic rats
Administration of Jiang-Zhi-Ning (a tradi- (Mukherjee et al. 1995c). The LD50 of the extract
tional Chinese medicine containing fleeceflower was found to be 2 g/kg. The extract (300 mg/kg
root, Fructus crataegi, Folium nelumbinis and and 600 mg/kg, orally) caused a reduction of
Semen cassiae) extract and effective fraction sig- blood glucose levels in streptozotocin-induced
nificantly reduced contents of serum total choles- diabetic rats by 53 % and 55 %, respectively, at
terol (TC), triglyceride (TG), low-density the end of 12 h.
lipoprotein-cholesterol (LDL-C), coronary index The crude protein of lotus seeds caused a
and atherogenic index in hyperlipidaemic rats as significant decrease (44.80 %) in the blood glu-
well as significantly increased contents of high- cose level of diabetic albino rats after 2 weeks
density lipoprotein-cholesterol (HDL-C), in rats of treatment (Ibrahim and El-Eraqy 1996). Oral
(Chen et al. 2011). Moreover, they significantly administration of lotus rhizome ethanolic
enhanced the activity of serum total nitric oxide extract significantly markedly reduced the
synthase (NOS) and increased contents of nitric blood sugar level of normal, glucose-fed hyper-
oxide. They also caused significant reductions in glycaemic and streptozotocin-induced diabetic
contents of endothelin-1 and malondialdehyde as rats, when compared with control animals
well as significant increase in superoxide dis- (Mukherjee et al. 1997b). The extract improved
mutase (SOD) activity and total antioxidant glucose tolerance and potentiated the action of
capacity (T-AOC) in the hyperlipidaemic rats. exogenously injected insulin in normal rats.
In-vitro, they restored and enhanced the vitality When compared with tolbutamide, the extract
of HUVECs with a concentration-dependent exhibited activity of 73 and 67 % of that of tol-
manner as well as content of NO in the culture butamide in normal and diabetic rats, respec-
media of HUVEC. They caused reductions in the tively. A methanol extract of the lotus stamens
contents of endothelin-1 and malondialdehyde in exerted an inhibitory effect on rat lens aldose
HUVECs. They also caused an increase in the reductase (RLAR) which had been shown to
vitality of SOD and T-AOC in HUVECs. play an important role in the complications asso-
Furthermore, they enhanced LDL-RmRNA ciated with diabetes (Lim et al. 2006). Among the
expression andCYP7A1 mRNA with in a isolated flavonoids, those harbouring 3-O-α-l-
concentration-dependent manner. Finally, they rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside
caused reductions in the contents of cholesterol groups in their C rings, including kaempferol
in Bel-7402. They also increased intercellular 3 - O - α - l - r h a m n o py r a n o s y l - ( 1 → 6 ) - β - d -
content of total bile acid as well as lowered extra- glucopyranoside and isorhamnetin 3-O-α-l-
cellular contents of TBA in the cells in a rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside,
concentration-dependent manner. Studies found exhibited the highest degree of rat lens aldose
84 Nelumbonaceae

reductase inhibitory activity in-vitro, with IC50 val- or symptoms of toxicity or mortality. The find-
ues of 5.6 and 9.0 μM, respectively. ings indicated that lotus flower extract had non-
Studies by Pan et al. (2009) demonstrated that acute toxicity but possessed hypoglycaemic and
neferine, from lotus embryo, had effects similar hypolipidaemic activities.
to rosiglitazone in significantly decreasing fast- Among several compounds extracted from
ing blood glucose, insulin, triglycerides and antidiabetic plants, nuciferine from N. nucifera
tumour necrosis factor-α (TNF-α) and enhancing was found to stimulate insulin secretion in iso-
insulin sensitivity in insulin-resistant rats. lated beta INS-1E cells by blocking potassium-
Streptozotocin-induced diabetic rats treated with adenosine triphosphate channels, thus
lotus seed ash at a concentration of 200 mg/kg contributing to the antidiabetic effects of
body weight orally for 30 days exhibited signifi- Nelumbo nucifera (Nguyen et al. 2012). Studies
cant hypoglycaemic activity (Mani et al. 2010). showed that lotus plumule polysaccharide (LPPS)
The hypoglycaemic activity of the ash was com- administration on non-obese diabetic (NOD)
parable with gliclazide. The presence of trace female mice for 15 weeks significantly increased
elements in appreciable amounts in the seeds pancreatic islet cell numbers and slightly
may play a direct or indirect role on insulin secre- enhanced the basal insulin secretion ability com-
tion or its action in a synergetic manner. Lotus pared to the control group (Liao and Lin 2013a).
leaf methanolic extract (NNE) increased insulin LPPS administration improved serum lipid pro-
secretion from β cells (HIT-T15) and human files in the diabetic mice via relatively increasing
islets in-vitro (Huang et al. 2011a). NNE serum high-density lipoprotein-cholesterol, but
enhanced the intracellular calcium levels in β decreasing low-density lipoprotein-cholesterol
cells and could also enhance phosphorylation of and total cholesterol levels. All four lotus leaf
extracellular signal-regulated protein kinases compounds, 2-hydroxy-1-methoxyaporphine,
(ERK)1/2 and protein kinase C (PKC). In-vivo pronuciferine, nuciferine and roemerine, showed
studies showed that NNE possessed the ability to the effects of improving insulin-stimulated glu-
regulate blood glucose levels in fasted normal cose consumption in differentiated 3T3-L1 adi-
mice and high-fat diet-induced diabetic mice. Of pocytes compared with the control group (Ma
its active constituents, quercetin did not affect et al. 2014). 2-hydroxy-1-methoxyaporphine and
insulin secretion, but catechin significantly and pronuciferine exhibited the most potent glucose
dose dependently enhanced insulin secretion. consumption-stimulatory activity at the concen-
Orally administered catechin significantly tration of 2 μg/mL.
reversed the glucose intolerance in high-fat diet-
induced diabetic mice. The findings suggested
that NNE and its active constituent catechin were Anti-inflammatory Activity
useful in the control of hyperglycaemia in non-
insulin-dependent diabetes mellitus through their Kaempferol 3-O-glucoside and luteolin
action as insulin secretagogues. Administration 7-O-glucoside from lotus aerial parts exhibited
of lotus flower extract at a dose of 250 mg/kg sig- anti-inflammatory activity (Wassel et al. 1996).
nificantly decreased the levels of fasting blood Methanol lotus rhizome extract at doses of 200
glucose (FBG), total cholesterol (TC), triglycer- and 400 mg/kg and betulinic acid at doses of car-
ides (TG), low-density lipoprotein (LDL) and rageenin and serotonin induced rat paw oedema
blood urea nitrogen (BUN) but increased high- in rats (Mukhjee et al. 1997a). The effects pro-
density lipoprotein (HDL) in streptozotocin- duced were comparable to that of phenylbuta-
induced diabetic rats compared to untreated zone and dexamethasone, two prototype
diabetic controls (Sakuljaitrong et al. 2013). anti-inflammatory drugs. Studies showed that
However, the extract did not alter creatinine in kaempferol, constituent of lotus extract, when
diabetic-treated rats compared to diabetic con- administered to rat tissues, attenuated the NF-κB
trols. Also the extract did not produce any signs nuclear binding activity (Kim et al. 2007).
Nelumbo nucifera 85

Kaempferol augmented GSH (glutathione) levels armepavine, 4′-methyl-N-methylcoclaurine,

in a dose-dependent manner and inhibited ROS apigenin-6-C-α- l -glucopyanosyl-8-C-β- d -
generation through the inhibition of iNOS and glucopyranoside and apigenin-6-C-α-l-
TNF-α expression in aged rat gingival tissues, via arabofuranosyl-8-C-β-d-glucopyranoside from
the modulation of the NF-κB and mitogen- lotus plumule; b-sitosterol from rhizome; nucif-
activated protein kinase (MAPK) pathways. erine, O-nornuciferine, hirsutrin, hyperin, luteo-
N. nucifera leaf extract demonstrated anti- loside and romaine from leaves; β-sitosterol from
inflammatory effects by inhibiting the production rhizome; and apigenin-6-C-α-l-arabofuranosyl-
of NO in RAW 264.7 cell activated by lipopoly- 8-C-β-d-glucopyranoside and an unknown com-
saccharide (Kwon et al. 2012). The inhibition of pound from seeds. Lotus leaf extract significantly
NO production was achieved by the inhibition of reduced the inflammatory effects of lipopolysac-
expression of iNOS protein. Additionally, the charide (LPS) in murine macrophage cell line
extract inhibited interleukin IL-6, one of cyto- RAW264.7 (Liu et al. 2014). Its active constitu-
kines related to inflammatory reaction. ents, quercetin and catechin, significantly also
Administration of methanolic seed extracts of red decreased the LPS-elevated protein expression of
and white lotus varieties at 400 mg/kg and iNOS, COX-2 and phospho-JNK. Besides, the
600 mg/kg to albino rats exerted anti- mRNAs and levels of IL-6 and TNF-α also
inflammatory effects in the carrageenin-induced decreased by quercetin and catechin treatment in
inflammation as well as in COX-2 enzyme inhi- LPS-induced RAW264.7 cells. The results
bition assay (Chakravarthi and Gopakumar showed that lotus leaf extract and its major com-
2010). Studies found that the purified compo- ponents quercetin and catechin exhibited anti-
nents, F1 and F2 from lotus plumule polysaccha- inflammatory activities by inhibiting the JNK- and
ride, exhibited potent anti-inflammatory effects NF-κB-regulated pathways and could therefore
on LPS-induced inflamed RAW264.7 macro- be a useful anti-inflammatory agent.
phages in a dose-dependent and preventive man-
ner (Liao and Lin 2012). Both F1 and F2,
especially F2, exhibited strong anti-inflammatory Central Nervous System (CNS)
effects in the presence of LPS by inhibiting toll- Activity (Sedative, Anxiolytic)
like receptor (TLR)-2 and/or TLR-4 expressions
in murine primary splenocytes in normal, inflam- Nuciferine exhibited depressant properties on the
matory and repair situations (Liao and Lin CNS in rodents, as well as weak anti-inflammatory,
2013b). The results further suggested that F2, a analgesic, antitussive, antiserotonin and adrener-
glycoprotein with low molecular weight of gic blocking activities (Macko et al. 1972). Lotus
25.7 kDa, may serve as a promising lead for the rhizome methanolic extract caused a reduction in
development of selective TLR antagonistic spontaneous activity; decreased exploratory
agents for inflammatory diseases. behavioural pattern by the head dip and Y-maze
Nuclear-factor-kappaB (NF-κB) inhibitors test; reduced muscle relaxant activity by rotarod,
with anti-inflammatory ability were identified in 30° inclined screen and traction test; and potenti-
various organs of lotus plant (Zhou et al. 2013a). ated the pentobarbitone-induced sleeping time in
Seventeen such compounds were found, ten mice significantly (Mukherjee et al. 1996c).
NF-κB inhibitors were found abundantly in lotus The methanol extracts of Nelumbo nucifera
embryo plumule (LP), most of which were iso- seed embryos significantly inhibited locomotor
quinoline alkaloids or flavone C-glycosides. LP activity in mice (Sugimoto et al. 2008). The chlo-
extracts were considered to have the best anti- roform partitioned extracts strongly inhibited
inflammatory ability compared to extracts from locomotor activity in mice, and its main alkaloid,
leaves and rhizome. The major compounds iden- neferine, dose dependently inhibited locomotor
tified were higenamine, lotusine, 4′-methylco- activity in mice. Neferine induced hypothermia
claurine, isoliensinine, liensinine, neferine, in mice and apparently potentiated thiopental-
86 Nelumbonaceae

induced sleeping time. An anxiolytic, diazepam lotus embryos, inhibited mice locomotor activity
decreased locomotor activity and rectal tempera- indicating that they all possessed sedative activ-
ture and enhanced sleep elicited by thiopental, ity (Nishimura et al. 2013). The efficacy of lien-
similar to neferine. In addition, neferine and sinine, isoliensinine and nelumboferine was more
diazepam showed anti-anxiety effects in the ele- potent than neferine and O-methylneferine.
vated plus maze test. Neferine did not affect mus- Compounds isolated from lotus leaves
cle coordination by the rotarod test. Neferine did 6′-hydroxy-4,4′-dimethoxychalcone, rhamnetin-
not affect strychnine- nor picrotoxin-induced sei- 3-O-β-d-glucopyranoside, quercetin-3-O-β-d-
zure. The results suggested that neferine had sev- glucopyranoside, quercetin-3-O-α- l-
eral central nervous system effects and that rhamnopyranosyl, quercetin-3-O-rutinoside and
neferine may participate in the efficacy of the isorhamnetin-3- O -α- l - rhamnopyranosyl-
sedative effects of embryos of N. nucifera seeds. (1 → 6)-[α- d - lyxopyranosyl-(1 → 2)-β- d -
Results of studies showed that lotus seed extract glucopyranoside] showed inhibitory activities
ameliorated scopolamine-induced amnesia in against beta amyloid (1–42) by A-beta aggrega-
rats by inhibiting acetylcholinesterase activity tion method with inhibition rates of (63.99)%,
and inducing choline acetyltransferase expres- (79.61)%, (49.96)%, (101.19)%, (88.41)% and
sion (Oh et al. 2009). The i.p. administration of (72.48)%, respectively (Zhao et al. 2013b). Of
neferine 25–100 mg/kg elicited anti-immobility the alkyl 4-hydroxybenzoates isolated from lotus
effects in mice (Sugimoto et al. 2010). The seeds, methyl 4-hydroxybenzoate and ethyl
molecular dose effects of neferine in the forced 4-hydroxybenzoate enhanced 5-HT-stimulated
swimming test were almost equal to those of the inward current (I5-HT) in Xenopus oocytes, but butyl
typical antidepressants maprotiline and imipra- 4-hydroxybenzoate reduced it (Youn et al. 2010).
mine. The antidepressant effect was found to be The results indicated that butyl 4-hydroxybenzoate
mediated by serotonin1A (5-HT1A) receptor and was an inhibitor of the 5-HT3A receptors
was likely to be linked to serotonergic expressed in Xenopus oocytes.
neurotransmission. N-methylasimilobine, a potent acetylcholines-
Compounds cycloartenol, p-hydroxybenzoic terase (AChE) from N. nucifera leaves, inhibited
acid, vanilloloside and nuciferoside, isolated 50 % of AChE activity at the concentrations of
from lotus stamens, exhibited good and noncom- 1.5 μg/mL when the IC50 value of physostigmine
petitive inhibition against acetylcholinesterase standard was 0.013 μg/mL (Yang et al. 2012).
(AChE) with IC50 values of 11.89, 20.07, 4.55 The mode of AChE inhibition by N-methyla-
and 3.20 microM and Ki values of 15.71, 25.44, similobine was reversible and noncompetitive.
7.76 and 5.76 μM, respectively (Jung et al.
2010b). Cycloartenol, p-hydroxybenzoic acid
and nuciferoside also possessed butyrylcholines- Cognitive-Enhancing Activity
terase (BChE) inhibitory activities with IC50 val-
ues of 13.93, 62.29, 205.78 and 83.06 μM, The methanol extract of N. nucifera rhizome elic-
respectively. The selectivity index (SI) values of ited in significant improvements of memory
1, 2, 3 and 5, calculated from IC50 values of BChE functions and neurogenesis in the dentate gyrus
and AChE, were 1.2, 3.1, 45.7 and 26.0. However, of Wistar rats (Yang et al. 2008). In the passive
all isolated compounds lacked BACE1 inhibition avoidance test, the retention time of extracted-
up to 100 μM. The data suggested that N. nucifera treated rats was significantly longer than that of
stamen-derived compounds could potentially controls. In addition, cell proliferation and cell
exert their primary anti-Alzheimer effects as differentiation increased significantly in the den-
AChE inhibitors rather than BACE1 inhibitors. tate gyrus. The results suggested that N. nucifera
Neferine, liensinine, isoliensinine, nelum- rhizome extract may improve learning and mem-
boferine and O-methylneferine, isolated from ory by enhancing neurogenesis in the dentate
Nelumbo nucifera 87

gyrus of the hippocampus. Hydroalcohol lotus increased the latency time measured during the
extract exhibited a weak inhibition of acetylcho- passive avoidance task in scopolamine-induced
linesterase with IC50value of 185.55 μg/mL, com- cholinergic blockade and normal naïve mice.
pared to physostigmine (IC50 0.076 μg/mL) used Allantoin treatment (3 or 10 mg/kg, for 7 days)
as a standard (Mukherjee et al. 2007). also increased the expression levels of phosphor-
Neferine, from lotus embryo, exerted signifi- ylated phosphatidylinositide 3-kinase (PI3K),
cant improvement in cognitive impairment in phosphorylated protein kinase B (Akt) and phos-
scopolamine-induced amnesia animal models phorylated glycogen synthase kinase-3β
and moderate inhibitory activities in cholinester- (GSK-3β). Also, allantoin significantly increased
ases (ChEs)- and β-site APP cleaving enzyme 1 the neuronal cell proliferation of immature neu-
(BACE1) assays (Jung et al. 2010a). Additionally, rons in the hippocampal dentate gyrus region.
it exhibited notable scavenging activities The results suggested that allantoin may have
against 1,1-diphenyl-2-picrylhydrazyl (DPPH), therapeutic potential for the cognitive dysfunc-
2,2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic tions observed in Alzheimer’s disease.
acid) diammonium salt (ABTS), NO (nitric
oxide) and O2− (superoxide) radicals, as well as
ONOO− (peroxynitrite). Neferine also demon- Convulsant Activity
strated dramatic inhibitory activity against lipid
peroxidation and protein nitration in cell-free Nornuciferine derivatives administered intraperi-
antioxidant assays and moderate inhibitory activ- toneally to mice as hydrobromide or hydrochlo-
ity of NO generation with exceptional suppres- ride provoked intense clonic convulsions
sion of NF-κB activation in cell-based assays. (Burkman and Cannon 1972). Unlike apomor-
The results demonstrated that the anti-amnesic phine, they did not induce emesis in dogs.
effect of neferine may be mediated via antioxi- N-Propyl nuciferine was the most effective con-
dant and anti-inflammatory capacities, as well as vulsant agent.
inhibition of ChEs and BACE1. The administra-
tion of lotus rhizome extract significantly amelio-
rated the scopolamine-induced reduction of Analgesic Activity
Ki67- and doublecortin-immunoreactive cells in
the hippocampal dentate gyrus of rats (Yoo et al. Kaempferol 3-O-glucoside and luteolin
2011). In addition, the administration of the 7-O-glucoside from lotus aerial parts exhibited
extract significantly restored the scopolamine- analgesic activity (Wassel et al. 1996). Methanol
induced reduction of brain-derived neurotrophic seed extract of red and white lotus, fed to albino
factor (BDNF) in hippocampal dentate gyrus rats, elicited considerable analgesic effect in the
homogenates. The results suggested that lotus acute pain (tail flick) model in a concentration-
rhizome extract could ameliorate the dependent fashion (Chakravarthi and Gopakumar
scopolamine-induced reductions of cell prolifer- 2009). The higher dose group of white lotus seed
ation, neuroblast differentiation and BDNF (600 mg/kg) exhibited more pronounced activity
levels. than other extracts.
Lotus plant extract was one of several plants
used in Indian Ayurvedic system of medicine for
improving cognitive function that showed IC50 Cosmeceutical/Antityrosinase
values <100 μg/ml for acetylcholinesterase Activity
inhibitory activity (Mathew and Subramanian
2014). Allantoin, an active compound in lotus, Lotus plant parts were found to have whitening
exerted memory-enhancing effects in mice (Ahn effect as measured by tyrosinase inhibition assay
et al. 2014). Subchronic administration of allan- and DOPA-oxidase inhibition assay, and anti-
toin (1, 3 or 10 mg/kg, for 7 days) significantly wrinkle effect as measured by elastase inhibition
88 Nelumbonaceae

assay (Kim et al. 2011). DOPA-oxidase inhibi- dose-dependent manners. These hepatoprotective
tion effect (whitening effect) of N. nucifera’s effects of lotus seed extract against CCl4 and
leaf, seed and flower extract was 59 %, 57 % and AFB1 might result from its potent antioxidative
50 %, respectively. N. nucifera’s leaf, seed and properties. In-vitro studies showed that neferine
flower extract showed 56 %, 49 % and 54 % elas- efficiently inhibited cultured hepatic stellate
tase inhibition (antiwrinkle effect) at 200 μg/ml, (HSC-T6) cell activation and induced apoptosis
while adenosine indicated 26 % inhibition. Lotus by increasing Bax and caspase 3 mRNAs expres-
plant parts were found to have whitening effect as sion via the mitochondrial pathway (Ding et al.
measured by tyrosinase inhibition assay and 2011).
DOPA-oxidase inhibition assay and antiwrinkle Administration of hydroalcoholic lotus seed
effect as measured by elastase inhibition assay extract to Wistar rats at 100 and 200 mg/kg body
(Kim et al. 2011). DOPA-oxidase inhibition weight for 4 days prior to carbon tetrachloride
effect (whitening effect) of N. nucifera’s leaf, treatment caused a significant dose-dependent
seed and flower extract was 59 %, 57 % and increase in the level of superoxide dismutase and
50 %, respectively. N. nucifera’s leaf, seed and catalase and a significant decrease in the level of
flower extract showed 56 %, 49 % and 54 % elas- thiobarbituric acid reactive substances (TBARS),
tase inhibition (antiwrinkle effect) at 200 μg/ml, when compared to CCl4-treated control in both
while adenosine indicated 26 % inhibition. Water liver and kidney (Rai et al. 2006). These changes
cream including Nelumbo nucifera’s root, leaf, observed at 100 mg/kg body weight treatment
flower and stem extract did not cause significant were comparable to those observed for standard
skin irritation. Methanolic extracts from the vitamin E at 50 mg/kg treatment. Armepavine, a
flower buds and leaves of sacred lotus were found lotus compound, exerted anti-hepatic fibrogenic
to show inhibitory effects on melanogenesis in effects in-vitro and in-vivo (Weng et al. 2009). It
theophylline-stimulated murine B16 melanoma dose dependently attenuated tumour necrosis
4A5 cells (Nakamura et al. 2013). Among the factor-α (TNF-α)- and lipopolysaccharide (LPS)-
constituents isolated, nuciferine, N-methyla- stimulated α-SMA protein expression and AP-1
similobine, (–)-lirinidine and 2-hydroxy-1-me- activation by HSC-T6 cells without adverse cyto-
thoxy-6a,7-dehydroaporphine showed potent toxicity in bile duct-ligated (BDL) rats. It signifi-
inhibition of melanogenesis. In addition, cantly reduced plasma AST and ALT levels,
3–30 μM nuciferine and N-methylasimilobine hepatic α-SMA expression and collagen contents
inhibited the expression of tyrosinase mRNA, and fibrosis scores as compared with vehicle
3–30 μM N-methylasimilobine inhibited the treatment. The results suggested that armepavine
expression of TRP-1 mRNA and 10–30 μM both in-vitro and in-vivo has antifibrotic effects
nuciferine inhibited the expression of TRP-2 in rats, possibly through anti-NF-κB activation
mRNA. pathways. Oral administration of lotus root hot
water extract with taurine supplementation
exerted antioxidant and hepatic protective effects
Hepatoprotective Activity in high-fat diet-induced obese male Sprague–
Dawley rats (Du et al. 2010). Activities of serum
Ethanol lotus seed extract showed potent DPPH glutamate oxaloacetate transaminase, glutamate
free radical scavenging effects with a median pyruvate transaminase and thiobarbituric acid
inhibition concentration of 6.49 g/ml (Sohn et al. reactive substance content were reduced, and
2003). Treatment of hepatocytes with the extract hepatic antioxidant enzymes were reduced by
inhibited both the production of serum enzymes lotus root extract. An ethanolic lotus leaf extract
and cytotoxicity by carbon tetrachloride (CCl4). at doses of 300 and 500 mg/kg exhibited hepato-
The genotoxic and cytotoxic effects of aflatoxin protective activity against CCl4-induced liver
B1 (AFB1) were also inhibited by the extract in toxicity in rats and antioxidant activity at 100 mg/
Nelumbo nucifera 89

kg that was comparable with that of a standard ameliorated high-fat diet-induced dyslipidaemia
treatment comprising 100 mg/kg of silymarin, a as well as liver steatosis and hepatic injury (Guo
known hepatoprotective drug (Huang et al. et al. 2013). The beneficial effects of nuciferine
2010b). were associated with altered expression of
Lotus germ oil treatment of mice with chronic hepatic genes involved in lipid metabolism. Tang
hepatotoxicity induced by carbon tetrachloride et al. (2014) showed that the polyphenol-rich
increased significantly the activities of antioxi- lotus leaf extract inhibited alcohol-induced ste-
dant enzymes, such as superoxide dismutase atohepatitis by multiple ways including reducing
(SOD), catalase (CAT) and the concentration of hepatic lipid synthesis, oxidative stress and anti-
glutathione (GSH) in a dose-dependent manner inflammation and enhancing fatty acid oxidation
(Lv et al. 2012). Also the lipid peroxidation prod- and transport responses in C57BL/6 J mice.
uct malondialdehyde and serum levels of hepatic
enzyme biomarkers (alanine aminotransferase
and aspartate aminotransferase) were decreased. Pulmonoprotective Activity
In addition, lotus germ oil could inhibit the con-
version of super-coiled pBR322 plasmid DNA to Administration of isoliensinine, a constituent of
the open circular form and apoptosis of hydrogen lotus embryo, markedly suppressed the increase
peroxide-induced PC-12 cells. in hydroxyproline content and abated pulmonary
After 3 weeks of feeding, the hepatomegaly fibrosis histological injury induced by bleomycin
and hepatic triglyceride accumulation was mark- in mice (Xiao et al. 2005). Isoliensinine enhanced
edly alleviated in the lotus polyphenol-diet-fed serum superoxide dismutase activity and
db/db mice relative to the control mice (Tsuruta decreased the bleomycin-elevated malondialde-
et al. 2011, 2012). Although the lipolytic enzyme hyde level in a concentration-dependent manner.
activity was not changed, the activities of lipo- Moreover, isoliensinine also significantly inhib-
genic enzymes, such as fatty acid synthase and ited the overexpression of TNF-α and TGF-β1
malic enzyme, were significantly lower in the induced by bleomycin. The results indicated that
lotus polyphenol-diet-fed db/db mice. Further, isoliensinine possessed a significant inhibitory
the presence of B-type proanthocyanidin poly- effect on bleomycin-induced pulmonary fibrosis,
mers with polymerisation degree up to 9 was probably due to its antioxidant and/or anti-
detected in the polyphenolic lotus root extract. It inflammatory activities and its inhibition of over-
was suggested that condensed tannins contained expression of TNF-α and TGF-β1 induced by
in lotus root could alleviate hepatic steatosis by bleomycin. Neferine, from lotus embryo, signifi-
suppressing the lipogenic enzyme activity in the cantly inhibited amiodarone-induced pulmonary
livers of db/db mice. They also found that the fibrosis in adult Kunming mice modulated by its
serum levels of adiponectin, which had been properties of anti-inflammation, surfactant pro-
reported to have a protective effect against non- tein-D inhibition and restoring increased
alcoholic fatty liver disease (NAFLD), were sig- CD4+CD25+ regulatory T cells which may regu-
nificantly higher in the Lotus group than in the late Th1/Th2 imbalance by suppressing Th2
control group of the db/db mice. Additionally, the response (Niu et al. 2013).
hepatic expression of such inflammatory genes as
tumour necrosis factor-alpha and monocyte che-
moattractant protein-1 was markedly suppressed Renoprotective Activity
by the Lotus diet. They speculated that the devel-
opment and progression of NAFLD were pre- Studies showed that (S)-armepavine, from lotus,
vented by suppressing the expression of lipogenic administered in the established phase of autoim-
and inflammatory genes as a result of the higher mune crescentic glomerulonephritis (ACGN) in
serum adiponectin level in the Lotus diet-fed db/ mice, was capable of markedly decreasing glo-
db mice. Nuciferine supplementation in hamsters merular crescents in the kidney and improving
90 Nelumbonaceae

proteinuria and renal dysfunction (Ka et al. action; its mechanism may be related to blockade
2010). These therapeutic effects were associated of Ca2+, Na+ influx (Wang et al. 1992). Liensinine
with (1) early systemic negative modulation of 3 mg/kg i.v. temporarily inhibited all parameters
T/B cells; (2) intra-renal regulation of combined of haemodynamics in anesthetised or pithed rats.
NF-κB activation and mononuclear leukocytic The dose-dependent inhibitory effects of liensi-
infiltration, thereby preventing glomerular cres- nine (1–30 mg/kg) on LVP, +dp/dtmax and SAP
cent formation; and (3) protection from apoptosis (slow action potential) in anesthetised rats were
in both the spleen and kidney. slightly stronger than those of quinidine 3 mg/kg.
Liensinine 1–30 mg/kg dose dependently pro-
duced these actions. The magnitude of inhibi-
Spasmolytic/Anti-arrhythmic/ tory effect of liensinine on all haemodynamic
Hypotensive/Bradycardic Activities parameters nearly corresponded to these of
verapamil 1 mg/kg. Liensinine 1–100 µmol/L
Nn-9 a non-crystalline alkaloid isolated from reduced the contractile force of isolated left
lotus embryo exerted hypotensive effects in ani- atria and the spontaneously beating rate of iso-
mals (Chen et al. 1962). In anesthetised cats, i.v. lated right atria of rabbits in concentration-
injections of 1–2 mg/kg lowered, after a delay of dependent manner. Liensinine 10–100 µmol/L
1/2–1 min, the blood pressure by about 50 % was also shown to concentration dependently
from the original level, lasting 2–3 h. In dogs, the decrease the action potential amplitude (APA)
blood pressure returned to normal in 1/2 h. This and the maximal velocity of phase 0 depolarisa-
dosage was ineffective in rabbits. Pretreatment tion (Vmax) and prolong the sinus cycle length
with Nn-9 also lessened the hypotensive action of (SCL) of SAP in isolated sinoatrial node pace-
histamine. Nn-9 appeared to act primarily maker cells of rabbits (Wang et al. 1993).
through histamine release. The alkaloids asimilo- Moreover, liensinine 1–100 µmol/L was found
bine and lirinidine, isolated from lotus leaves, to inhibit the slow inward current of canine car-
inhibited the contraction of isolated rabbit aorta diac Purkinje fibre in concentration-dependent
induced by serotonin; the pA2 values were 5.78 manner. Liensinine 3 and 100 µmol/L reduced
and 7.36, respectively (Shoji et al. 1987). the peak value of slow inward current by 14 %
Studies found that neferine from lotus embryo and 88 %, respectively. The results suggested
possessed antiarrhythmic action (Li et al. 1989). that liensinine possessed calcium antagonistic
It exerted an inhibitory effect on the slow trans- effects.
membrane Na+ and/or Ca2+ current of myocar- Studies by Morales et al. (1997) demonstrated
dium. Neferine decreased the amplitude of action the hypotensive and bradycardic properties of
potential and the maximal upstroke velocity in (±)-norarmepavine, a lotus alkaloid. In rat’s aorta,
rabbit sinoatrial nodes and in the clusters of cul- (±)-norarmepavine (10 mg/kg i.v.) decreased the
tured cardiac myocytes from neonatal rats in a mean arterial pressure and heart rate by 45 % and
concentration-dependent manner. In subsequent 21 %, respectively. It showed a negative chrono-
studies, neferine at 1–10 mg/kg i.v. dose depend- tropic effect on rat-isolated atria, decreasing the
ently decreased the monophasic action potential spontaneous frequency by about 54 %. Aortic
amplitude, prolonging the monophasic action rings contracted with KCl 70 mM were relaxed
potential duration in anaesthetised cat’s heart (Li in a concentration-dependent manner by
et al. 1990). It also decreased left ventricular (±)-norarmepavine, (±)-coclaurine and (±)-norco-
pressure (LVP), dP/dt, prolonged sinus cycle claurine. The two earlier alkaloids exhibited an
length and reduced arterial blood pressure in a efficacy similar to verapamil, relaxing the aortic
dose-dependent manner. These effects were simi- rings by 100 %. (±)-Norcoclaurine exhibited a
lar to those of quinidine and different from tet- lower efficacy. The rank order of potency was
randrine. Liensinine, an alkaloid from lotus verapamil > (±)-norarmepavine > (±)-norcoclau-
embryo, was shown to have antiarrhythmic rine > (±)-coclaurine.
Nelumbo nucifera 91

Neferine alkaloid isolated from lotus embryo aggregation is related to regulation of TXA2/
exhibited antiarrhythmic effect in experimental PGI2 and cAMP/cGMP balance. Studies found
arrhythmic models (Qian 2002). In guinea pig neferine could dose dependently inhibit human
papillary muscles and atria, neferine reduced the or rat platelet aggregation induced by ADP, col-
force of contraction, decreased the amplitude and lagen and epinephrine both in-vitro and in-vivo
upstroke velocity Vmax of action potential and (Qian 2002). Neferine, from lotus embryo, sig-
prolonged action potential duration (APD50 and nificantly and dose dependently inhibited colla-
APD 90) and effective refractory period. Neferine gen-, thrombin- and U46619-induced platelet
also suppressed the contractility of rabbit papil- aggregation in mice-washed platelets, or ADP-
lary muscles, prolonged functional refractory induced platelet aggregation in mice platelet-rich
period and inhibited automaticity, positive stair- plasma (Zhou et al. 2013b). Neferine treatment
case and post-rest potentiation. Neferine, isolated decreased platelet dense granule secretion initi-
from the seed embryo, increased the basal cAMP ated by collagen, thrombin and U46619. Also,
concentration in rabbit corpus cavernosum tissue Neferine markedly and dose dependently pro-
in a dose-dependent manner (Chen et al. 2008). moted the dissociation of platelet aggregates pre-
The accumulation of cAMP induced by prosta- formed by various agonists including collagen,
glandin E1 (PGE1, a stimulator of cAMP produc- thrombin, U46619 or ADP. Neferine could sig-
tion) was also augmented by neferine in a nificantly reduce the area of mice platelet adhe-
dose-dependent manner. It was found that nefer- sion to the collagen and inhibited thrombosis
ine with its spasmolytic activity could enhance in-vitro. In collagen–epinephrine-induced acute
the concentration of cAMP in rabbit corpus cav- pulmonary thrombus mouse model, neferine, at
ernosum tissue, probably by inhibiting phospho- 6 mg/kg, significantly attenuated thrombus
diesterase activity. Neferine, from lotus embryo, formation.
induced a concentration-dependent decrease in
current amplitude according to the voltage steps
and tail currents of human ether-à-go-go-related Immunomodulatory Activity
gene (HERG) K+ channels in a dose-dependent
manner with an IC50 of 7.419 μM (Gu et al. 2009) N. nucifera ethanolic extracts significantly sup-
in human embryonic kidney (HEK293) cells. It pressed phytohemagglutinin-activated human
had no effect on the generation and trafficking of peripheral blood mononuclear cell (PBMC) pro-
HERG protein. A blocked-off HERG channel liferation (Liu et al. 2004). The inhibitory action
was one mechanism of the antiarrhythmic effects of lotus extract did not involve direct cytotoxic-
by neferine. ity. The suppressant effects of lotus extract on
proliferation of activated PBMC appeared to be
mediated, at least in part, through inhibition of
Antithrombotic/Antiplatelet Activity early transcripts of PBMC, especially those of
important interleukin IL-2, interferon-gamma
Neferine was shown to significantly inhibit rabbit (IFN-gamma) and cyclin-dependent kinase (cdk)
platelet aggregation induced by ADP, collagen, 4 mRNA expression and arrest of cell cycle pro-
arachidonic acid (AA) and platelet-activating gression in the cells. Oral administration of MRL/
factor (PAF) with IC50 of 16, 22, 193 and MpJ-lpr/lpr mice with similar disease features to
103 μmol/L, respectively (Yu and Hu 1997). human systemic lupus erythematosus with
Neferine was found to increase vascular 6-keto- (S)-armepavine (from lotus) for 6 weeks pre-
PGF1 alpha and platelet cAMP levels in dose- vented lymphadenopathy and elongated life span
dependent manner, but inhibit AA-stimulated of MRL/MpJ-lpr/lpr mice (Liu et al. 2006). It
TXA2 release from platelets. The results sug- appeared to be mediated by inhibition of spleno-
gested that the mechanism of neferine on platelet cyte proliferation, suppression of interleukin-2
92 Nelumbonaceae

(IL-2), interleukin-4, interleukin-10, and Cardiovascular Protective Activity

interferon-gamma (IFN-gamma) gene expres-
sions, reduction of glomerular hypercellularity Isoliensinine, extracted from lotus embryo,
and immune complexes deposition and decrease inhibited angiotensin II-induced proliferation of
of urinary protein and anti-double-stranded DNA porcine coronary arterial smooth muscle cells
autoantibody production. It was suggested that in-vitro (Xiao et al. 2006). Its antiproliferative
(S)-armepavine may be an immunomodulator for effect was related to the decrease of the overex-
the management of autoimmune diseases like sys- pression of growth factor platelet-derived
temic lupus erythematosus. Also, they found that growth factor (PDGF)-β and basic fibroblast
(S)-armepavine inhibited phytohemagglutinin- growth factor (bFGF), proto-oncogene c-fos,
activated human peripheral blood mononuclear c-myc and hsp70. Polyphenol-rich water extract
cell (PBMC) proliferation and cytokine produc- of lotus leaves was found to inhibit both prolif-
tion in a major way by blocking membrane- eration and migration of vascular smooth mus-
proximal effectors such as IL-2-inducible T-cell cle cells (VSMC) and thus may serve as a
kinase (Itk) and phospholipase Cgamma potential anti-atherogenic agent (Ho et al.
(PLCgamma) in a PI-3K-dependent manner 2010). 1.0 % of lotus leaf extract reduced neo-
(Liu et al. 2007). intima formation conspicuously and inhibited
Studies found that the extract of lotus rhizome smooth muscle cell proliferation and MMP-2
and seed stimulates defence system by modu- secretion in the blood vessel of rabbits fed with
lating several immunological parameters a high-cholesterol diet. Neferine, from lotus
(Mukherjee et al. 2010b). Total leukocyte count embryo, inhibited nitric oxide production
and lymphocyte count increased significantly, but induced by lysophosphatidylcholine in human
the neutrophil count was decreased in both umbilical vein endothelial cells (HUVECs) by
extract-treated animal groups compared to the modulating dimethylarginine dimethylamino-
control. A dose-dependent potentiation of hydrolase (DDAH) – asymmetric dimethylargi-
delayed type hypersensitivity reaction induced nine (ADMA) pathway via its antioxidant
by sheep red blood cells was observed from the properties (Peng et al. 2011). Animal studies
extracts. The percentage of neutrophil adhesion showed that administration of lotus leaf extract
to the nylon fibre was increased in rhizome- elicited cardioprotective effect in isoproterenol
treated groups (63.22 and 62.91 %) compared to (ISO)-induced myocardial infarction in rats
the seed-treated group (54.86 and 54.23 %). A (Subashini et al. 2012). Oral pretreatment with
potential phagocytic response was seen on treat- the extract (400 mg/kg) to ISO-induced rats
ment of the extracts, and significant changes daily for a period of 21 days showed a signifi-
were observed in the formation of formazone cant decrease in the levels of lipid peroxidative
crystals. In separate studies, lotus rhizome extract products and improved the antioxidant enzyme
stabilised erythrocyte membrane significantly at activities. This amelioration was supported by
10 (42.05 %) and 100 μg/mL (44.31 %) histopathological findings.
(Mukherjee et al. 2010a). The extract showed Oral administration of lotus leaf extract for
38.66 % (100 μg/mL) and 69.66 % (10 μg/mL) 4 weeks to rats after balloon injury reduced the
degranulation against compound 48/80 (C 48/80) intimal thickening by suppressing vascular
in mast cells. The extract at 1 and 5 μg/mL inhib- smooth muscle cells (VSMC)’s proliferation
ited lipopolysaccharide (LPS)-induced activation through inhibition of extracellular signal-
of macrophages by decreasing the expression of regulated kinase 1/2 phosphorylation and their
costimulatory molecules. The extracts also inhib- migration by reducing the expression of MMP-2
ited the nitrite concentration at 1 and 5 μg/mL and MMP-9 through inhibition of JNK1/2 phos-
compared to LPS-treated group. phorylation (Karki et al. 2013). The results
Nelumbo nucifera 93

suggested that lotus leaf could be considered to temperature (pyrexia) in rats (Mukherjee et al.
have therapeutic value in the prevention of 1996a). The effect produced was comparable
atherosclerosis. with the standard antipyretic drug, paracetamol
(150 mg/kg, i.p.). The ethanol lotus stalk extract
at a dose of 200 mg/kg was found to produce sig-
Aldose Reductase Inhibitory Activity nificant lowering of normal body temperature up
to 3 h, and at 400 mg/kg, it caused significant
Among several solvent fractions of lotus leaf lowering of body temperature up to 6 h after its
extract, the ethyl acetate and n-butanol fractions, administration (Sinha et al. 2000). In the model
having prominent total phenolic content (TPC) of yeast-induced elevation of body temperature,
and total flavonoid content (TFC)C values, the extract showed dose-dependent lowering of
showed significant antioxidant effects in the body temperature up to 4 h at both the doses, and
DPPH and TEAC assays (Jung et al. 2008). the results were comparable to that of paracetamol,
Moreover, the ethyl acetate fraction exhibited a standard antipyretic agent.
superior inhibitory effects in the total reactive
oxygen species (ROS), rat lens aldose reductase
(RLAR) and advanced glycation end products Antihemorrhagic Activity
(AGE) assays, with IC50 values of 9.4, 2.4 and
28.2 μg/ml, respectively. Also, two of its key The antihemorrhagic principle in nelumbins
antioxidant flavonoids, quercetin 3-O-β-d- receptaculum, dried receptacle of Nelumbo
glucopyranoside and quercetin 3-O-β-d- nucifera, was identified as quercetin [2-(3, 4-dihy-
glucuronopyranoside, may play important roles droxy)-3, 5, 7-trihydroxy-4H-1-benzopyran-4-
in the antioxidant and RLAR inhibitory effects of one] (Ishida et al. 1988).
N. nucifera leaves. The hydroalcoholic extract of
lotus plant exhibited aldose reductase inhibitory
activity with an IC50 value 59.78 μg/mL, and its Antiallergic Activity
alkaloidal extract was more inhibitory with an
IC50 of 28.82 μg/mL (Gupta et al. 2014). Flow cytometric analysis revealed that Fc epsilon
RI expression on human basophilic KU812F
human basophilic KU812F cell surface was sup-
Diuretic Activity pressed in a concentration-dependent manner
when the cells were cultured with kaempferol
The methanol extract of lotus rhizomes induced extracted from a methanolic extract of flavonoid-
significant diuresis in rats (Mukherjee et al. rich lotus stamens (Shim et al. 2009). Moreover,
1996b). Dose-dependent effects were observed in RTPCR analysis showed that the mRNA levels for
urine volume and electrolyte excretion. There was Fc epsilon RI alpha- and gamma-chains were
a significant increase in natriuretic and chlorouretic reduced as the result of kaempferol treatment in a
activity, but kaliuresis was less than natriuresis. concentration-dependent manner. Kaempferol
showed its suppressive effects on intracellular Ca2+
concentration and histamine release from anti-Fc
Antipyretic Activity epsilon RI alpha-chain antibody-stimulated cells
in a concentration-dependent manner. These
The methanolic lotus rhizome extract, in doses of observations indicated that kaempferol may exert
200, 300 or 400 mg/kg (p.o.), produced signifi- antiallergic effect via downregulation of Fc epsi-
cant dose-dependent lowering of normal body lon RI expression and degranulation.
temperature and yeast-induced elevation of body
94 Nelumbonaceae

Antiestrogenic/Antisteroidogenic doses (100, 200, 400 and 600 mg/kg body wt.)
Activity reduced not only the frequency of defecation,
wetness of faecal dropping and PGE2-induced
The petroleum ether lotus seed extract adminis- enteropooling but also the propulsive movements
tered to mice exhibited antiestrogenic, antipro- of charcoal meal significantly in rats (Talukder
gestational and contraceptive activity at a dose of and Nessa 1998). Aqueous lotus fruit extract was
3 mg/kg body weight (Mazumder et al. 1992). found to have strong significant anti-rotaviral
Oral administration of lotus seed petroleum ether activity with a 50 % inhibitory concentration
extract to sexually immature female rats and (IC50) <300 μg/ml (Knipping et al. 2012). A com-
mature male rats on alternate day for 15 days bination of Glycyrrhiza glabra and lotus showed
caused a remarkable delay in sexual maturation synergy in their antiviral activities and interven-
in prepubertal female rats as evidenced from age tions of lotus, and combination may be useful in
of vaginal opening and first estrus (cornified the treatment of diarrhoea caused by rotavirus
smear) and a significant reduction in the sperm infection.
count and motility in mature male rats (Gupta
et al. 1996). In both the cases, treatment resulted
in accumulation of cholesterol and ascorbic acid Anti-dermatitis Activity
and reduction in D5-3b-hydroxysteroid dehydro-
genase and glucose-6-phosphate dehydrogenase Oral administration of lotus leaf extract to
activity in the ovary and testis of female and male NC/Nga mice with atopic dermatitis induced
rat, respectively. The results indicated suppres- by repeated epicutaneous application of
sion of steroidogenesis in both ovary and testis by 2,4-dinitrochlorobenzene (DNCB) on the dorsal
lotus seed extract. skin resulted in the suppression of clinical sever-
Oral administration of lotus seed ethanol ity score, transepidermal water loss, scratching
extract to female albino rats elicited in significant behaviour and blood IgE level (Karki et al. 2012).
decline in weights of reproductive organs (ovary, Histopathologic analyses revealed that thicken-
uterus, vagina) (Mutreja et al. 2008). Glycogen ing of the epidermis and mast cell degranulation
levels in the reproductive organs were decreased were significantly reduced in lotus-treated mice.
significantly, but the cholesterol levels were The results suggested that lotus leaf extract may
increased; no changes in haematological param- be a useful natural resource for the management
eters were found. The extract prolonged the oes- of atopic dermatitis.
trous cycle. The results suggested that lotus seed
exhibited antiestrogenic activity without altering
the general physiology of the female rats. Antimalarial Activity

Antimalarial activity was observed for lotus leaf

Antidiarrhoeal Activity alkaloids (R)-roemerine and N-methylasimilobine
with IC50 = 0.2 and 4.8 μg/mL for the D6
Rats treated with lotus rhizome methanol extract (chloroquine-sensitive) clone of Plasmodium fal-
significantly inhibited castor oil-induced diarrhoea ciparum, respectively, and 0.4 and 4.8 μg/mL for
and PGE-induced enteropooling (Mukherjee the W2 (chloroquine-resistant) clone, respec-
et al. 1995b). It also showed significant reduction tively (Agnihotri et al. 2008). An analysis of the
in gastrointestinal motility following charcoal structure–activity relationship revealed that the
meal in rats. The results obtained confirmed the substituents in position C-1 and C-2 of aporphine
efficacy of lotus rhizome extract as an antidiar- alkaloids were crucial for the antimalarial
rhoeal agent. Lotus rhizome extract in graded activity.
Nelumbo nucifera 95

Radioprotective/Photoprotective Candida albicans, Aspergillus niger, Aspergillus

Activity fumigatus and Trichophyton mentagrophytes
(Mukherjee et al. 1995d). Antibacterial activity
Pretreatment of Swiss albino mice with lotus seed in-vitro was documented for lotus rhizome
pod procyanidins (LSPCs) 200 mg/kg by intragas- extracts against Staphylococcus aureus,
tric (i.g.) for 15 days before gamma irradiation Escherichia coli, Bacillus subtilis, Bacillus pumi-
was found to be the most effective dose in prevent- lus and Pseudomonas aeruginosa (Mukherjee
ing radiation sickness, reducing radiation-induced et al. 1995a). Kaempferol 3-O-glucoside and
mortality, increasing mean survival time and ele- luteolin 7-O-glucoside from lotus aerial parts
vating radiation median lethal dose (LD50) from showed significant activity against six bacteria
8.9 to 10.5 Gy, indicating a dose-modifying factor and Candida albicans (Wassel et al. 1996). Lotus
(DMF) of 1.18 (Duan et al. 2010). Further, admin- pollen essential oil significantly inhibited growth
istered LSPCs at a dose of 200 mg/kg could effec- of Salmonella typhimurium and Escherichia coli
tively maintain spleen index close to normal; in-vitro (Sittiwet 2009). The MICs and MBCs
stimulate endogenous spleen colony forming were 10–40 and 20–80 ml/L, respectively. The
units; promote the levels of red blood cells (RBC), hydroethanolic extract of both white and pink
white blood cells (WBC), platelets and haemoglo- lotus flowers showed almost similar antimicro-
bin in peripheral blood and prevent spleen and skin bial activities (Brindha and Arthi 2010a). MIC
damage in irradiated mice; reduce the level of for white flower extract against Escherichia coli
radiation-induced micronucleated polychromatic and Staphylococcus aureus was found to be
erythrocytes in bone marrow, maintain the poly- 430 μg and 450 μg, respectively, and pink flower
chromatic erythrocytes (PCE) and normochro- showed 480 μg and 490 μg, respectively.
matic erythrocytes (NCE) ratio (P/N ratio); and The pink flower extract showed moderate in-
significantly decrease bone marrow chromosomal vitro inhibitory activity against Aspergillus
damage. Alternatively, pretreatment with LSPCs niger and Monascus purpureus, while the white
(200 mg/kg) significantly decreased the lipid per- flower extract had weaker activity. Hexane
oxidation (LPO) level and elevated the activities of extract of lotus seed oil strongly inhibited
endogenous antioxidant enzymes in liver after growth in-vitro of diarrheal bacteria Shigella
irradiation. The results indicated that LSPCs pos- sp., Staphylococcus aureus and Klebsiella sp.
sessed potent whole body radioprotective activity, and in a moderate way Salmonella sp and
and it may be used as a radioprotector. weakly against Escherichia coli (Anitha and
Lotus leaf extract (10, 100 and 1000 μg/ml) Arunkumar 2012). It also inhibited fungal der-
possessed strong protective effect against UVB- matophytes Malassezia furfur, Trichophyton
induced phototoxicity in the mitochondria model mentagrophytes and T. rubrum.
(Huang et al. 2013). The in-vivo test showed that
LLE lotus leaf extract exerted significant protec-
tive effects on the level of superoxide dismutase, Antiosteoporotic Activity
catalase and glutathione peroxidase, as well as
the contents of hydroxyproline and malondialde- Im et al. (2013) found that water and ethanol
hyde in mouse skin. lotus seed extracts protected L6 rat skeletal mus-
cle cells from antimycin-induced mitochondria-
mediated cell death. The extracts reduced cellular
Antimicrobial Activity apoptosis; preserved the mitochondrial mem-
brane potential; protected mitochondrial ATP
The methanol lotus rhizome extract inhibited the production; inhibited p53, Bax, and caspase 3
in-vitro growth of the following fungi-tested activities; and induced Bcl-2 production.
96 Nelumbonaceae

Prebiotic Activity 1.10 ppm; LC90 = 28.13, 35.83 and 3.59 ppm),
respectively. Of nine plant extracts tested ,the
Studies showed that prebiotics such as lotus seed- highest larvicidal activity was observed after
resistant starch, especially P-LRS3, could stimu- 24 h in the leaf methanol extract of N. nucifera,
late the growth of Bifidobacterium adolescentis seed ethyl acetate and methanol extract of P.
(Zhang et al. 2013a). Compared with glucose and nigrum against the fourth instar larvae of
high amylose maize starch (HAMS) media, bifi- Anopheles stephensi (LC50 = 34.76, 24.54 and
dobacteria had a higher tolerance to gastrointesti- 30.20 ppm) and against Culex quinquefasciatus
nal conditions in lotus-resistant starches LRS3 (LC50 = 37.49, 43.94 and 57.39 ppm), respec-
and P-LRS3 media. The rough surface of lotus- tively (Kamaraj et al. 2011). Lotus leaf ethyl
resistant starch and the short chain fatty acids acetate extract caused 100 % mortality of A. ste-
produced during fermentation might influence phensi and C. quinquefasciatus after 48 h expo-
the proliferation of bifidobacteria. Lotus seed- sure. The maximum repellent activity was
resistant starch was more effective than either observed at 500 ppm in methanol extracts of N.
glucose or HAMS in promoting the proliferation nucifera, ethyl acetate and methanol extract of P.
of bifidobacteria (Zhang et al. 2014). nigrum and methanol extract of Trachyspermum
ammi. Lotus leaf extracts were found to have
insecticidal (adulticidal) activity (Santhoshkumar
Anthelmintic Activity et al. 2011). In adults of Haemaphysalis bispi-
nosa, parasitic mortality of 80, 74, 72 and 100 %
Aporphine alkaloids from lotus leaves, liriode- was observed in lotus crude leaf hexane, ethyl
nine, lysicamine, (–)-anonaine, (–)-asimilobine, acetate, acetone and methanol extracts, respec-
(–)-cadaverine, (–)-N-methylasimilobine, tively. Maximum efficacy was observed in lotus
(–)-nuciferine, (–)-nornuciferine, (–)-roemerine, leaf methanol extract against the adults of the
7-hydroxydehydronuciferine and cepharadione B Acarina H. bispinosa and the hematophagous fly
killed Hymenolepis nana or reduced their sponta- Hippobosca maculata with LC50 and LC90 values
neous movements (oscillation/peristalsis) but had of 437.14 and 200.81 ppm, respectively.
no larvicidal effect or ability to halt spontaneous
parasite movement for 72 h of Anisakis simplex
(Lin et al. 2014). The above compounds showed Drug Interaction/Cytochrome
greater lethal anthelmintic efficacy on H. nana Inhibitory Activity
than against A. simplex.
Studies showed that nuciferine inhibited cyto-
chrome P450 1A2 (CYP1A2) activity in-vitro
Insecticidal/Larvicidal Activity and caused changes in the pharmacokinetic
parameters of phenacetin after oral administra-
The hexane, chloroform, ethyl acetate, acetone, tion to Wistar rats (Hu et al. 2010). Studies
methanol and aqueous lotus leaf extracts and syn- revealed that the alcoholic lotus leaf extract and
thesised silver nanoparticles using aqueous lotus its alkaloid fraction strongly inhibited human
leaf extract showed moderate larvicidal effects cytochrome CYP2D6 with an IC50 value of
against fourth instar larvae of Anopheles subpic- 12.05 μg/mL and 0.966 μg/mL, respectively, and
tus and Culex quinquefasciatus (Santhoshkumar weakly inhibited other isoenzymes (CYP2C9,
et al. 2011). Maximum efficacy was observed in CYP2C19, CYP2E1 and CYP3A4) (Ye et al.
crude methanol, aqueous and synthesised silver 2014a). The flavonoid fraction was found to
nanoparticles against the larvae of A. subpictus weakly inhibit CYP2D6. The three aporphine
(LC50 = 8.89, 11.82 and 0.69 ppm; LC90 = 28.65, alkaloids isolated from the alkaloid fraction,
36.06 and 2.15 ppm) and against the larvae of C. nuciferine, N-nornuciferine and 2-hydroxy-1-
quinquefasciatus (LC50 = 9.51, 13.65 and methoxyaporphine significantly inhibited
Nelumbo nucifera 97

CYP2D6 with IC50 values of 3.78, 3.76 and uronic acid and 2-hydroxy-1-methoxyaporphine-
3.15 μM, respectively. 2-O-sulphate. Following a single oral and
intravenous administration of total bisbenzyliso-
quinoline alkaloids from lotus seed embryo in
Mutagenic/Antimutagenic Activity rats, liensinine, isoliensinine and neferine were
detected in rat plasma (Huang et al. 2011b). The
The nitrite-treated methanol extract of Nelumbo lower limit of quantification can reach 0.03 μg/ml
nucifera flowers exhibited the high mutagenicity for the three compounds. Studies found that
on Salmonella typhimurium TA 98 and TA 100 neferine had no effect on amiodarone plasma
strains (Wongwattanasathien et al. 2010). All pharmacokinetics when it was co-administered
dichloromethane extracts of flowers including intragastrically with amiodarone, an antiarrhyth-
lotus decreased the mutagenicity induced by the mic agent (Wan et al. 2011). Studies in Caco-2
product of 1-aminopyrene nitrite model on both cells demonstrated that liensinine, neferine and
tester strains. isoliensinine, from lotus embryo, were substrates
of P-glycoprotein, whereas MRP2 cell was not
involved in the transport process, suggesting that
Toxicity Studies P-glycoprotein might be responsible for the
absorption and distribution of the 3 alkaloids (Yu
None of the lotus leaf alkaloid compounds were et al. 2013b).
cytotoxic to Vero cells up to a concentration of After administration of neferine to rats, one-
23.8 μg/mL (Agnihotri et al. 2008). The results compartmental pharmacokinetic parameters indi-
from both acute and subchronic oral toxicity cated that nuciferine had rapid distribution,
studies suggested that the oral lethal dose of lotus extensive tissue uptake and poor absorption into
stamen extract for male and female rats was in systemic circulation (Gu et al. 2014). The values
excess of 5000 mg/kg, and the no-observed- of absolute bioavailability were (3.8)%, (4.2)%
adverse-effect level (NOAEL) of the extract for and (3.9)% after oral administration of 2.0, 5.0
both male and female rats was considered to be and 10.0 mg/kg nuciferine and intravenous
200 mg/kg/day (Kunanusorn et al. 2011). administration of 0.2 mg/kg nuciferine in rats.
The results suggested that nuciferine was distrib-
uted into the brain, liver and adipose tissue after
Pharmacokinetic Studies intravenous administration. A total of 37 in-vivo
components were identified in rats after oral
Nornuciferine was found to be metabolised by rat administration of a lotus leaf flavonoid extract,
and rabbit liver microsomes to lysicamine (Smith including quercetin-3-O-glucuronide, quercetin-
and Sood 1971). Rat, rabbit and guinea pig liver 3-O-glucoside, quercetin-3-O-galacoside, quer-
microsomes also dealkylated N-alkylated analog cetin and kaempferol, as well as their methylation,
of nornuciferine. 2-Hydroxy-1-methoxyaporphin glucuronidation and sulfonation metabolites (Ye
(2H1M), an alkaloid from Nelumbo nucifera, one et al. 2014b).
of the herbal component of the Chinese
Traditional Medicine Jiang-Zhi-Ning (JZN), was
identified as the constituent showing the major
pharmacodynamic effect (Chen et al. 2011, Traditional Medicinal Uses
2012a). The major metabolites of 2H1M after
intragastric administration of JZN to beagle dogs Nelumbo nucifera is a medical plant used in tra-
were analysed and identified as N-demethyl-2- ditional Chinese medicine, and every part is uti-
hydroxy-1-methoxyaporphine-2-O-glycuronic lised and officially listed in the Chinese
acid, 2-hydroxy-1-methoxyaporphine-2-O-glyc- Pharmacopoeia as:
98 Nelumbonaceae

– Lianzi, Semen Nelumbinis, the dried mature (Nelumbo nucifera Gaertn) roots have been used
seeds used as a sedative and tonic. as an antidiabetic and an antiproliferative remedy
– Lianzixin, Plumula Nelumbinis, the dried (Park et al. 2009).
embryos from the mature seeds used as a sed- In folk medicines, seeds are used in the treat-
ative and haemostatic. ment of tissue inflammation, cancer, skin dis-
– Lianfang, Receptaculum Nelumbinis, the eases and leprosy; used as poison antidote,
dried receptacles used as a haemostatic. antiemetic and cooling agent; and generally pre-
– Lianxu, Stamen Nelumbinis, the dry stamens scribed to children as diuretic and refrigerant
collected when the lotus flowers bloom used (Chopra et al. 1986; Liu et al. 2004). The fruits
as an adstringent. and seeds of lotus are astringent and used to treat
– Heye, Folium Nelumbinis, the dry leaves used hyperdipsia, dermatopathy, halitosis, menorrha-
as a haemostatic. gia, leprosy and fever (Nadkarni and Nadkarni
– Oujie, Nodus Nelumbinis rhizomatis, the dry 1982). Seed powder mixed with honey is useful
nodes of lotus rootstock used as a haemostatic in treating cough, while roots with ghee (melted
(Tang and Eisenbrand 1992). After charring fresh butter), milk and gold promote strength,
the receptacles, leaves and rootstocks are also virility and intellect. Its rhizomes are used for
used against bleeding. According to Chinese treating pharyngopathy, pectoralgia, spermator-
traditional knowledge, medicinal uses of dif- rhoea, leucoderma, small pox, diarrhoea, dysen-
ferent part of lotus plants are common in the tery and cough (Mukherjee et al. 1997a, b; Ou
treatment of diarrhoea, tissue inflammation 1989). The stem is used in indigenous Ayurvedic
and haemostasis (Yu and Hu 1997). medicines as a diuretic and anthelmintic and to
treat strangury, vomiting, leprosy, skin diseases
Nelumbo nucifera leaf has been used for sum- and nervous exhaustion. Young leaves with sugar
mer heat syndrome as home remedy in Japan and are useful to treat rectal prolapse, and the leaves
China, and it has recently been used to treat obe- boiled with Mimosa pudica in goat’s milk can be
sity in China (Ono et al. 2006). Embryo of lotus used to treat diarrhoea. Nelumbo nucifera has
seeds is used in traditional Chinese drug called been used for the treatment of several disorders
‘Lian Zi Xin’, which primarily helps to overcome including skin disease, cough, inflammation,
nervous disorders, insomnia, high favours (with fever and many other disorders (Mukherjee et al.
restlessness) and cardiovascular diseases (e.g. 2010b; Ou 1989). The leaves are used as an effec-
hypertension, arrhythmia) (Chen et al. 2007). tive drug for haematemesis, epistaxis, haemoptysis,
Jiang-Zhi-Ning (JZN), a TCM, is composed of haematuria, metrorrhagia, fever and inflamma-
four Chinese herbs, i.e. fleeceflower root, Fructus tory skin conditions (Ou 1989).
Crataegi, Folium Nelumbinis and Semen Cassiae Nelumbo nucifera seed has been used as an
is used to strengthen blood circulation of coro- antiobesity agent in traditional Chinese herbal
nary artery, arrhythmia and hyperlipidaemia medicine (Pan et al. 2009). Hyperlipidaemia in
(Chen et al. 2011). Lotus leaf is known to be rodents can be treated with lotus leaves (La Cour
effective for ‘overcoming body heat’ and stop- et al. 1995; Onishi et al. 1984). Leaves also pos-
ping bleeding and is commonly used as a tradi- sess diuretic and astringent properties and help to
tional curing plant for the treatment of treat fever, sweating and strangury and as styptic
haematemesis, epistaxis, haemoptysis, haematu- (Chinese Materia Medica 1977).
ria and metrorrhagia in traditional Chinese medi- The Chinese administer the rhizomes for diar-
cine (Lee and Lee 2011). The herb of lotus rhoea and dysentery; the Cambodians make a tea
(Nelumbo nucifera) leaves is the traditional for menorrhagia (Burkill 1966). The Chinese
Chinese medicine He Ye, which is commonly used the embryo to reduce high fever and in the
used to treat sunstroke, assuage thirst and cure treatment of cholera, haemoptysis and spermator-
both diarrhoea and fever in China (Ye et al. rhoea. In Malayan medicine, it is highly esteemed
2014a, b). In traditional Korean medicine, lotus as a tonic and taken as tea for fever. The Malays
Nelumbo nucifera 99

used pounded petals for syphilis. They are astrin- vested 4–5 months after planting the lotus.
gent and taken for diarrhoea and vomiting in Additionally, lotus shoots provide substrate for
Java. The Chinese used them in facial cosmetic. growth of epiphytic algae that are consumed by
In India, they are used for fever and in the tilapia fish. Lotus rhizomes knot, as a waste from
Philippines for dysentery. The stamens are astrin- food production, will be a potential material for
gent and diuretic and used by the Chinese. They extracting antioxidants (Hu and Skibsted 2007).
are also used in cosmetics. In India, they are In India, the fibres from the stem are used as lamp
employed as an astringent and cooling medicine. wicks (Burkill 1966). A Thai cigarette may be
In Indonesia and India, the juice from the petioles wrapped in lotus petal to impart a dainty
and peduncles is used for treating diarrhoea. appearance.
In Vietnam, the seeds are used to treat dysen- Studies by Thongtha et al. (2014) found that
tery, spermatorrhoea, leucorrhoea, palpitation, phosphorus in domestic wastewater was removed
insomnia, general debility and anorexia (NIMM from the initial concentration of 1.038 mg/L to
1999). The plumules are useful in the therapy of 0.094 and 0.048 mg/L by N. nucifera and Cyperus
permanent mild fever, insomnia, thirst and hae- alternifolius, respectively, within 5 days.
moptysis. The leaves are beneficial in treating Domestic wastewater is a source of phosphorus
bloody stools, haematuria, gingival and subcuta- contamination that causes eutrophication when it
neous haemorrhage. The stamens are effective contaminates aquatic environments.
for metrorrhagia, haemoptysis, spermotorrhoea Lotus leaf extract inhibited algal bloom more
and insomnia. The achnes in decoction are used effectively than the stem (rhizome) extract (He
for treating dysentery and aphonia. The recepta- et al. 2013). At 25 g/L, the leaf extract exhibited
cle in decoction is useful as haemostatic in treat- highest inhibition rate of 71.33 % and 88.14 %
ing bloody stools, urination, leucorrhoea and against the algae Microcystis aeruginosa and
hypertension. Rhizome decoction is employed as Scenedesmus quadricauda, respectively, while
haemostatic in treating bloody stools, haematu- the inhibition rate for the rhizome extract was
ria, haematemesis, epistaxis and metrorrhagia. 49.78 % and 52.14 %, respectively.
NIMM (1999) also reported that in India, a nutri-
tious flour made from the rhizome is given to
children affected by diarrhoea, dysentery and Comments
dyspepsia. A paste made of powdered rhizome is
applied for tinea and other dermatoses. Lotus is revered as a sacred religious flower in
Hinduism and Buddhism. Lotus flower is the
national flower of India and is legendary in
Other Uses folklore and religious mythology. In Japan,
Moriyama City’s prefectural flower is the lotus.
The lotus is a very beautiful and attractive plant The lotus flower is the national flower of Vietnam
cultivated in many parts of the world as an orna- and the golden lotus logo of Vietnam’s Airline.
mental and as cut flower used as centrepieces for
parties or as religious offerings in India and
Thailand. The dried stalked seed receptacle is Selected References
also used in dried floral arrangements. In India,
the fibre of the stem is used for making the wicks Agnihotri VK, ElSohly HN, Khan SI, Jacob MR, Joshi
VC, Smillie T, Khan IK, Walker LA (2008)
of sacred lamps.
Constituents of Nelumbo nucifera leaves and their
It is commonly planted in fields or ponds with antimalarial and antifungal activity. Phytochem Lett
nutrient-rich mud and can extract nutrients from 1(2):89–93
old pond mud efficiently. Water levels of ponds Ahn JH, Kim ES, Lee C, Kim S, Cho SH, Hwang BY, Lee
MK (2013) Chemical constituents from Nelumbo
can be increased as lotus plant grows. Fish can be
nucifera leaves and their anti-obesity effects. Bioorg
stocked when water levels reach 30 cm and har- Med Chem Lett 23(12):3604–3608
100 Nelumbonaceae

Ahn YJ, Park SJ, Woo H, Lee HE, Kim HJ, Kwon G, Gao Chaudhuri PK, Singh D (2009) A new lipid and other con-
Q, Jang DS, Ryu JH (2014) Effects of allantoin on stituents from the rhizomes of Nelumbo nucifera.
cognitive function and hippocampal neurogenesis. J Asian Nat Prod Res 11(7):583–587
Food Chem Toxicol 64:210–216 Chaudhuri PK, Singh D (2013) A new triterpenoid from
Anitha A, Arunkumar D (2012) Fatty acid composition the rhizomes of Nelumbo nucifera. Nat Prod Res
and antidermatophytic and antidiarrheal activity of 27(6):532–536
Nelumbo nucifera seed oil. Int J Pharm Pharm Sci Chen WZ, Chu CC, Ting KS (1962) Hypotensive mecha-
4(4):264–270 nism of the alkaloid NN-9 from Nelumbo nucifera.
Arthur HR, Cheung HT (1959) An aporphine alkaloid, Yao Xue Xue Bao 13:271–276 (In Chinese)
nuciferine, from Asiatic lotus cultivated in Hong Chen Y, Fan GR, Wu HL, Wu YT, Mitchell A (2007)
Kong. Chem Abstr 53(20):19306g Separation, identification and rapid determination of
Backer CA, Bakhuizen van den Brink RC Jr (1963) Flora liensine, isoliensinine and neferine from embryo of the
of Java, vol 1. Noordhoff, Groningen, 648 pp seed of Nelumbo nucifera Gaertn. by liquid chroma-
Bernauer K (1963) Pronuciferin, ein benzylisochinolin- tography coupled to diode array detector and tandem
alkaloid mit para-cyclohexadienon gruppierung. Helv mass spectrometry. J Pharm Biomed Anal 43:99–104
Chim Acta 46(5):1783–1785 (In German) Chen J, Liu JH, Wang T, Xiao HJ, Yin CP, Yang J (2008)
Bernauer K (1964) Uber die isolierung von (+)-pronucif- Effects of plant extract neferine on cyclic adenosine
erin und (–)-anonain aus den keimlingen von Nelumbo monophosphate and cyclic guanosine monophosphate
nucifera Gaertn. Helv Chim Acta 47(8):2119–2122 levels in rabbit corpus cavernosum in vitro. Asian
(In German) J Androl 10(2):307–312
Borsch T, Barthlott W (1996) Classification and distribu- Chen JX, Zhao HH, Yang Y, Liu B, Ni J, Wang W (2011)
tion of the genus Nelumbo Adans. (Nelumbonaceae). Lipid-lowering and antioxidant activities of Jiang-Zhi-
Beitr Biol Pflanzen 68:421–450 Ning in Traditional Chinese Medicine. J Ethnopharmacol
Boustie J, Stigliani JL, Montanha J, Amoros M, Payard 134(3):919–930
M, Girre L (1998) Antipoliovirus structure-activity Chen JX, Ma XL, Gao K, Wang Y, Zhao HH, Wu H, Wang
relationships of some aporphine alkaloids. J Nat Prod J, Xie H, OuYang YL, Luo LT, Guo SZ, Han J, Liu B,
61(4):480–484 Wang W (2012a) The active ingredients of Jiang-Zhi-
Brindha D, Arthi D (2010a) Antimicrobial activity of Ning: study of the Nelumbo nucifera alkaloids and
white and pink Nelumbo nucifera Gaertn flowers. their main bioactive metabolites. Molecules
J Pharm Res Health Care 2(2):147–155 17(8):9855–9867
Brindha D, Arthi D (2010b) Evaluation of Antitumor and Chen S, Fang LC, Xi HF, Guan L, Fang JB, Liu YL, Wu
in vivo antioxidant potentials of Nelumbo nucifera BH, Li SH (2012b) Simultaneous qualitative assess-
Gaertn (white and pink) flowers in Ehrlich Ascites ment and quantitative analysis of flavonoids in various
Carcinoma mice. J Pharm Res 3(10):2483–2487 tissues of lotus (Nelumbo nucifera) using high perfor-
Burkill IH (1966) A dictionary of the economic products mance liquid chromatography coupled with triple
of the Malay Peninsula. Revised reprint, 2 vol. quad mass spectrometry. Anal Chim Acta
Ministry of agriculture and co-operatives, Kuala 724:127–135
Lumpur, Malaysia. Vol 1 (A–H) pp 1–1240, Vol 2 Chen S, Wu BH, Fang JB, Liu YL, Zhang HH, Fang LC,
(I–Z) pp 1241–2444 Guan L, Li SH (2012c) Analysis of flavonoids from
Burkman AM, Cannon JG (1972) Screening nornucifer- lotus (Nelumbo nucifera) leaves using high perfor-
ine derivatives for apomorphine-like activity. J Pharm mance liquid chromatography/photodiode array detec-
Sci 61(5):813–814 tor tandem electrospray ionization mass spectrometry
Cai CH, Cai JW, Man JM, Yang Y, Wang ZF, Wei CX and an extraction method optimized by orthogonal
(2014) Allomorph distribution and granule structure design. J Chromatogr A 1227:145–153
of lotus rhizome C-type starch during gelatinization. Chen S, Xiang Y, Deng J, Liu YL, Li SH (2013)
Food Chem 142:408–415 Simultaneous analysis of anthocyanin and non-
Centre for Indigenous Peoples’ Nutrition and Environment anthocyanin flavonoid in various tissues of different
(CINE) (2007) Asian case study – Miao Sichuan lotus (Nelumbo) cultivars by HPLC-DAD-ESI-MSn.
China. Community food system data tables. McGill PLoS One 8(4):1–13
University, Canada. Chinese Pharmacopoeia Committee (2010)
resources/data/miao/ Pharmacopoeia of the People’s Republic of China, vol
Chakravarthi PV, Gopakumar N (2009) Evaluation of 1. Chemical Industry Press, Beijing, pp 193–195
analgesic activity of lotus seeds (Nelumbo nucifera) in Chopra RN, Nayar SL, Chopra IC (1986) Glossary of
albino rats. Vet World 2(9):355–357 Indian medicinal plants. (Including the Supplement).
Chakravarthi PV, Gopakumar N (2010) Anti-inflammatory Council Scientific Industrial Research, New Delhi,
activity of red and white lotus seeds (Nelumbo 330 pp
nucifera) in albino rats. Vet World 3(4):157–159 Chu P, Chen HH, Zhou YL, Li Y, Ding Y, Jiang LW,
Chao YC, Chou YL, Young PT, Chou TQ (1962) Studies Tsang EWT, Wu KQ, Huang SZ (2012) Proteomic and
on the alkaloids of embryo of loti Nelumbo nucifera. functional analyses of Nelumbo nucifera annexins
I. Isolation and characterization of liensinine. Sci involved in seed thermotolerance and germination
China Math 5(2):215–219 (In Chinese) vigor. Planta 235(6):1271–1288
Nelumbo nucifera 101

Council of Scientific and Industrial Research (CSIR) Fu D, Wiersema JH (2001) Nelumbonaceae dumortier. In:
(1966) The wealth of India. A dictionary of Indian raw Wu ZY, Raven PH, Hong DY (eds) Flora of China, vol
materials and industrial products. (Raw Materials 7). 6 (Caryophyllaceae through Lardizabalaceae). Science
Publications and Information Directorate, New Delhi, Press, Beijing, and Missouri Botanical Garden Press,
India St. Louis
Deng SG, Deng ZY, Fan YW, Li J, Liu R, Xiong DM Furukawa H (1965a) On the alkaloids of Nucifera nucifera
(2009a) Application of high-speed counter-current Gaertn. IX. Alkaloids of loti embryo. (2). Structure of
chromatography coupled with high performance liq- neferine, a new biscoclaurine alkaloid. Yakugaku
uid chromatography for the separation and purification Zasshi 85(4):335–338 (In Japanese)
of Quercetin-3-O-sambubioside from the leaves of Furukawa H (1965b) On the alkaloids of Nelumbo
Nelumbo nucifera. Front Chem Eng Chin 3:375–382 nucifera Gaertn. X. Alkaloids of loti embryo. (3).
Deng SG, Deng ZY, Fan YW, Peng Y, Li J, Xiong DM, Domestic “lien tze hsin”. Yakugaku Zasshi 85(4):353–
Liu R (2009b) Isolation and purification of three flavo- 354 (In Japanese)
noid glycosides from the leaves of Nelumbo nucifera Furukawa H (1966a) On the alkaloids of Nelumbo
(Lotus) by high-speed counter-current chromatogra- nucifera Gaertn. XII. Alkaloids of loti embryo.
phy. J Chromatogr B Analyt Technol Biomed Life Sci Yakugaku Zasshi 86(1):75–77 (In Japanese)
877(24):2487–2492 Furukawa H (1966b) Studies on the alkaloids of Nelumbo
Deng J, Chen S, Yin XJ, Wang K, Liu YL, Li SH, Yang PF nucifera Gaertn. NMR spectra of liensinine type alka-
(2013) Systematic qualitative and quantitative assess- loids. Yakugaku Zasshi 86(10):883–886 (In Japanese)
ment of anthocyanins, flavones and flavonols in the Furukawa H (1971) Studies on the alkaloids of Nelumbo
petals of 108 lotus (Nelumbo nucifera) cultivars. Food nucifera Gaertn.: NMR spectra of liensinine type alka-
Chem 139(1–4):307–312 loids. Yakugaku Zasshi 86(10):883–886 (In Japanese)
Ding YF, Cheng HY, Song SQ (2008) Changes in extreme Furukawa H, Yang TH, Lin TJ (1965) On the alkaloids of
high-temperature tolerance and activities of antioxi- Nelumbo nucifera Gaertn. XI. Alkaloids of loti
dant enzymes of sacred lotus seeds. Sci China C Life embryo. 4. Structure of lotusine, a new water-soluble
Sci 51(9):842–853 (In Chinese) quaternary base. Yakugaku Zasshi 85(5):472–475 (In
Ding H, Shi JH, Wang Y, Guo J, Zhao JH, Dong L (2011) Japanese)
Neferine inhibits cultured hepatic stellate cell activa- Gayathri K, Dhevi R, Niraimathi KL, Shabi MM,
tion and facilitates apoptosis: a possible molecular Subashini U, Rajamanickam GV, Dubhey GP (2009)
mechanism. Eur J Pharmacol 650(1):163–169 Comparative evaluation of Nelumbo nucifera (Gaertn.)
Do TC, Nguyen TD, Tran H, Stuppner H, Ganzera M flowers and vitamin C in In vitro antioxidant activity.
(2012) Quantitative determination of phenolic com- ICFAI J Life Sci 3(1):7–19
pounds in lotus (Nelumbo nucifera) leaves by capillary Goo HR, Choi JS, Na DH (2009) Simultaneous determi-
zone electrophoresis. Planta Med 78(16):1796–1799 nation of quercetin and its glycosides from the leaves
Do TCMV, Nguyen TD, Tran H, Stuppner H, Ganzera M of Nelumbo nucifera by reversed-phase high-
(2013) Analysis of alkaloids in lotus (Nelumbo performance liquid chromatography. Arch Pharm Res
nucifera Gaertn.) leaves by non-aqueous capillary 32(2):201–206
electrophoresis using ultraviolet and mass spectromet- Gu DF, Li XL, Qi ZP, Shi SS, Hu MQ, Liu DM, She CB,
ric detection. J Chromatogr A 1302:174–180 Lv YJ, Li BX, Yang BF (2009) Blockade of HERG K+
Du H, You JS, Zhao X, Park JY, Kim SH, Chang KJ channel by isoquinoline alkaloid neferine in the stable
(2010) Antiobesity and hypolipidemic effects of lotus transfected HEK293 cells. Naunyn Schmiedebergs
leaf hot water extract with taurine supplementation in Arch Pharmacol 380(2):143–151
rats fed a high fat diet. J Biomed Sci 17(1):S42–S47 Gu SY, Zhu GH, Wang YZ, Li Q, Wu X, Zhang JG, Liu
Duan Y, Zhang H, Xie B, Yan Y, Li J, Xu F, Qin Y (2010) GL, Li XY (2014) A sensitive liquid chromatography–
Whole body radioprotective activity of an acetone- tandem mass spectrometry method for pharmacoki-
water extract from the seedpod of Nelumbo nucifera netics and tissue distribution of nuciferine in rats.
Gaertn. Seedpod. Food Chem Toxicol J Chromatogr B 961:20–28
48(12):3374–3384 Guo H (2009) Cultivation of lotus (Nelumbo nucifera
Duan XH, Pei L, Jiang JQ (2013) Cytotoxic alkaloids Gaertn. ssp. nucifera) and its utilization in China.
from stems of Nelumbo nucifera. Zhongguo Zhong Genet Resour Crop Evol 56:323–330
Yao Za Zhi 38(23):4104–4108 (In Chinese) Guo MD, Chen LG (1984) Studies on the alkaloids con-
Duanmu QP, Li AF, Sun AL, Liu RM, Li XP (2010) Semi- stituents of the embryo nelumbinis (Nelumbo nucifera)
preparative high-speed counter-current chromatogra- produced in China. Chin Tradit Herb Drugs 15:291–
phy separation of alkaloids from embryo of the seed of 293 (In Chinese)
Nelumbo nucifera Gaertn by pH-gradient elution. Guo XF, Wang DJ, Duan WJ, Du JH, Wang X (2010)
J Sep Sci 33(12):1746–1751 Preparative isolation and purification of four flavo-
ENVIS (2008) Medicinal plants of conservation concern. noids from the petals of Nelumbo nucifera by high-
Environmental information System (ENVIS) Centre, speed counter-current chromatography. Phytochem
Foundation for Revitalisation of Local Health Anal 21(3):268–272
Traditions (FRLHT), Ministry of Environment and Guo FC, Yang X, Li XX, Feng RN, Guan CM, Wang YW,
Forests, India. Li Y, Sun CH (2013) Nuciferine prevents hepatic ste-
102 Nelumbonaceae

atosis and injury induced by a high-fat diet in ham- Huang CF, Chen YW, Yang CY, Lin HY, Way TD, Chiang
sters. PLoS One 8(5):e63770 W, Liu SH (2011a) Extract of lotus leaf (Nelumbo
Gupta M, Mazumder UK, Mukhopadhyay RK, Sarkar S nucifera) and its active constituent catechin with insu-
(1996) Antisteroidogenic effect of the seed extract of lin secretagogue activity. J Agric Food Chem
Nelumbo nucifera in the testis and the ovary of the rat. 59(4):1087–1094
Indian J Pharm Sci 58:236–242 Huang Y, Zhao LB, Bai Y, Liu P, Wang JL, Xiang JZ
Gupta S, Singh N, Jaggi AS (2014) Evaluation of in vitro (2011b) Simultaneous determination of liensinine,
aldose reductase inhibitory potential of alkaloidal isoliensinine and neferine from seed embryo of
fractions of Piper nigrum, Murraya koenigii, Nelumbo nucifera Gaertn. in rat plasma by a rapid
Argemone mexicana, and Nelumbo nucifera. J Basic HPLC method and its application to a pharmacoki-
Clin Physiol Pharmacol 25(2):255–265 netic study. Arzneimittelforschung 61(6):347–352
Ha JY, Lee K, Park J, Dong A, Shin HS (2010) Huang B, Zhu L, Liu S, Li D, Chen YX, Ma BX, Wang
Cytoprotective activity of lotus (Nelumbo nucifera YW (2013) In vitro and in vivo evaluation of inhibi-
Gaertner) leaf extracts on the mouse embryonic fibro- tion activity of lotus (Nelumbo nucifera Gaertn.)
blast cell. Food Sci Biotechnol 19(5):1171–1176 leaves against ultraviolet B-induced phototoxicity.
Han X, Wang W, Wu CA, Liu S, Wang LY (2008) Effect J Photochem Photobiol B 121:1–5
of nuciferine on cholesterol metabolism in Bel-7402 Hutadilok-Towatana N, Chaiyamutti P, Panthong K,
cells. Progr Modern Biomed 9:1628–1630 (In Mahabusarakam W, Rukachaisirikul V (2006)
Chinese) Antioxidative and free radical scavenging activities of
He LS, Meng FL, Diao XJ, Li YW, Meng R, Xi BD, Shu some plants used in Thai folk medicine. Pharm Biol
JM (2013) Allelopathic effect of Nelumbo nucifera 44(3):221–228
stem and leaf tissue extract on the growth of Hyun SK, Jung YJ, Chung HY, Jung HA, Choi JS (2006)
Microcystis aeruginosa and Scenedesmus quadri- Isorhamnetin glycosides with free radical and ONOO-
canda. Huan Jing Ke Xue 34(7):2637–2641 (In scavenging activities from the stamens of Nelumbo
Chinese) nucifera. Arch Pharm Res 29(4):287–292
Ho HH, Hsu LS, Chan KC, Chen HM, Wu CH, Wang CJ Ibrahim N, El-Eraqy W (1996) Protein content and amino
(2010) Extract from the leaf of nucifera reduced the acid composition of Nelumbo nucifera seeds and its
development of atherosclerosis via inhibition of vas- evaluation as hypoglycaemic agent. Egypt J Pharm Sci
cular smooth muscle cell proliferation and migration. 37(1-6):635–664
Food Chem Toxicol 48(1):159–168 Im AR, Kim YH, Uddin MR, Chae SW, Lee HW, Jung
Hsieh YY, Chen WC, Kao YS (1964a) Studies on the WS, Kim YH, Kang BJ, Kim YS, Lee MY (2013)
alkaloids of Nelumbo nucifera. III. Absolute configu- Protection from antimycin A-induced mitochondrial
ration of liensinine. Sci China Math 13:2018–2019 (In dysfunction by Nelumbo nucifera seed extracts.
Chinese) Environ Toxicol Pharmacol 36(1):19–29
Hsieh YY, Pan PC, Chen WC, Kao YS (1964b) Alkaloids Ishida H, Umino T, Tsuji K, Kosuge T (1988) Studies on
of Nelumbo nucifera. IV. Total synthesis of liensinine. the antihemorrhagic substances in herbs classified as
Sci Sin 12(12):2020–2025 (In Chinese) hemostatics in Chinese medicine. VIII On the anti-
Hu SY (2005) Food plants of China. The Chinese hemorrhagic principle in Nelumbins Receptaculum.
University Press, Hong Kong, 844 pp Chem Pharm Bull 36(11):4585–4587
Hu M, Skibsted LH (2007) Antioxidative capacity of rhi- Itoh A, Saitoh T, Tani K, Uchigaki M, Sugimoto Y,
zome extract and rhizome knot extract of edible lotus Yamada J, Nakajima H, Ohshiro H, Sun SJ, Tanahashi
(Nelumbo nuficera). Food Chem 76:327–333 T (2011) Bisbenzylisoquinoline alkaloids from
Hu XM, Zhou BH, Luo SD, Cai HS, Yin WH (1993) Nelumbo nucifera. Chem Pharm Bull 59(8):947–951
Quantitative determination of liensinine in the embryo Jeong IY, Lee HY, Park YD, Jin CH, Choi DS, Byun MW
Nelumbinis (Nelumbo nucifera Gaertn.) by TLC- (2009) Effects of gamma irradiation on total polyphe-
scanning. Zhongguo Zhong Yao Za Zhi 18(3):167– nols, radical scavenging activities and decolourization
168, 192. (In Chinese) of Nelumbo nucifera extracts. Radiat Phys Chem
Hu LW, Xu W, Zhang X, Su J, Liu XR, Li HY, Zhang WD 78:575–577
(2010) In-vitro and in-vivo evaluations of cytochrome Jiang JZ, Cao BS (2005) Cultivar types of aquatic vegeta-
P450 1A2 interactions with nuciferine. J Pharm bles and their utilization. Chin Food Nutr 9:21–24
Pharmacol 62(5):658–662 Jiang Y, Ng TB, Wang CR, Li N, Wen TY, Qiao WT,
Huang B, Ban XQ, He JS, Tong J, Tian J, Wang YW Zhang D, Cheng ZH, Liu F (2010) First isolation of
(2010a) Comparative analysis of essential oil compo- tryptophan from edible lotus (Nelumbo nucifera
nents and antioxidant activity of extracts of Nelumbo Gaertn) rhizomes and demonstration of its antioxidant
nucifera from various areas of China. J Agric Food effects. Int J Food Sci Nutr 61(4):346–356
Chem 58(1):441–448 Jung HA, Kim JE, Chung HY, Choi JS (2003) Antioxidant
Huang B, Ban XQ, He JS, Tong J, Tian J, Wang YW principles of Nelumbo nucifera stamens. Arch Pharm
(2010b) Hepatoprotective and antioxidant activity of Res 26(4):279–285
ethanolic extracts of edible lotus (Nelumbo nucifera Jung HA, Jung YJ, Yoon NY, da Jeong M, Bae HJ, Kim
Gaertn.) leaves. Food Chem 120(3):873–878 DW, Na DH, Choi JS (2008) Inhibitory effects of
Nelumbo nucifera 103

Nelumbo nucifera leaves on rat lens aldose reductase, whitening and anti-wrinkle cosmetic agent. Korean
advanced glycation endproducts formation, and oxida- J Chem Eng 28(2):424–427
tive stress. Food Chem Toxicol 46(12):3818–3826 Knipping K, Garssen J, van’t Land B (2012) An evalua-
Jung HA, Jin SE, Choi RJ, Kim DH, Kim YS, Ryu JH, tion of the inhibitory effects against rotavirus infection
Kim DW, Son YK, Park JJ, Choi JS (2010a) Anti- of edible plant extracts. Virol J 9(1):137–144
amnesic activity of neferine with antioxidant and anti- Koch K, Dommisse A, Barthlott W (2006) Chemistry and
inflammatory capacities, as well as inhibition of ChEs crystal growth of plant wax tubules of lotus (Nelumbo
and BACE1. Life Sci 87(13-14):420–430 nucifera) and nasturtium (Tropaeolum majus) leaves
Jung HA, Jung YJ, Hyun SK, Min BS, Kim DW, Jung JH, on technical substrates. Cryst Growth Des
Choi JS (2010b) Selective cholinesterase inhibitory 6(11):2571–2578
activities of a new monoterpene diglycoside and other Koshiyama H, Ohkuma H, Kawaguchi H, Hsu HY, Chen
constituents from Nelumbo nucifera stamens. Biol YP (1970) Isolation of 1-(p-Hydroxybenzyl)-6, 7-dihy-
Pharm Bull 33(2):267–272 droxy-1, 2, 3, 4-tetrahydroisoquinoline (demethylco-
Ka SM, Kuo YC, Ho PJ, Tsai PY, Hsu YJ, Tsai WJ, Lin claurine), an active alkaloid from Nelumbo nucifera.
YL, Shen CC, Chen A (2010) (S)-armepavine from Chem Pharm Bull 18(12):2564–2568
Chinese medicine improves experimental autoimmune Kredy HM, Huang DH, Xie BJ, He H, Yang ER, Tian BQ,
crescentic glomerulonephritis. Rheumatology Xiao D (2010) Flavonols of lotus (Nelumbo nucifera,
(Oxford) 49(10):1840–1851 Gaertn.) seed epicarp and their antioxidant potential.
Kamaraj C, Rahuman AA, Bagavan A, Elango G, Zahir Eur Food Res Technol 231:387–394
AA, Santhoshkumar T (2011) Larvicidal and repellent Kunanusorn P, Panthong A, Pittayanurak P,
activity of medicinal plant extracts from Eastern Ghats Wanauppathamkul S, Nathasaen N, Reutrakul V
of South India against malaria and filariasis vectors. (2011) Acute and subchronic oral toxicity studies of
Asian Pac J Trop Med 4(9):698–705 Nelumbo nucifera stamens extract in rats.
Karki R, Jung MA, Kim KJ, Kim DW (2012) Inhibitory J Ethnopharmacol 134(3):789–795
effect of Nelumbo nucifera (Gaertn.) on the development Kunimoto J, Kamimura M (1964) Studies on the alkaloids
of atopic dermatitis-like skin lesions in NC/Nga mice. of Nelumbo nuciefera Gaertn. VII. Cleavage reaction
Evid Based Complement Alternat Med 2012:153568 of nuciferine and nornuciferine by metallic sodium in
Karki R, Jeon ER, Kim DW (2013) Nelumbo nucifera leaf liquid ammonia. Yakugaku Zasshi 84(11):1100–1104
extract inhibits neointimal hyperplasia through modu- Kunitomo J, Yamamoto C, Otsuki T (1964) Studies on the
lation of smooth muscle cell proliferation and migra- alkaloids of Nelumbo nucifera Gaertn. VI. Alkaloids
tion. Nutrition 29(1):268–275 of “gen-shi-hasu”. Yakugaku Zasshi 84(11):1141–
Kashiwada Y, Aoshima A, Ikeshiro Y, Chen YP, Furukawa 1142 (In Japanese)
H, Itoigawa M, Fujioka T, Mihashi K, Cosentino ML, Kunitomo J, Nagai Y, Okamoto Y, Furukawa H (1970) On
Morris-Natschke SL, Lee KH (2005) Anti-HIV ben- the alkaloids of Nelumbo nucifera Gaertn. XIV. On the
zylisoquinoline alkaloids and flavonoids from the tertiary base. Yakugaku Zasshi 90(9):1165–1169 (In
leaves of Nelumbo nucifera and structure-activity cor- Japanese)
relations with related alkaloids. Bioorg Med Chem Kunitomo J, Miyoshi M, Maruyama T, Yuge E (1971)
13(2):443–448 Studies on the alkaloids of Nelumbo nucifera Gaertn.
Khattak KF, Simpson TJ, Ihasnullah (2009) Effect of XV. Synthesis of dl-nornuciferine (dl-1-methoxy-2-
gamma irradiation on the microbial load, nutrient hydroxyaporphine). Yakugaku Zasshi 91(8):896–900
composition and free radical scavenging activity of (In Japanese)
Nelumbo nucifera rhizome. Radiat Phys Chem Kunitomo J, Yoshikawa Y, Tanaka S, Imori Y, Isoi K,
78:206–212 Masada Y, Hashimoto K, Inoue T (1973) Alkaloids of
Kim MJ, Shin HS (2012) Antioxidative effect of lotus Nelumbo nucifera. Phytochemistry 12(3):699–701
seed and seedpod extracts. Food Sci Biotechnol Kuo YC, Lin YL, Liu CP, Tsai WJ (2005) Herpes sim-
21(6):1761–1766 plex virus type 1 propagation in HeLa cells inter-
Kim SB, Rho SB, Rhyu DY, Kim DW (2005) Effect of rupted by Nelumbo nucifera. J Biomed Sci
Nelumbo nucifera leaves on hyperlipidemic athero- 12(6):1021–1034
sclerotic Bio F1B hamster. Korean J Pharmacogn Kwon K, Kwon OY, Kang KH (2012) Anti-inflammatory
36(3):229–234 (In Korean) effects of Nelumbo nucifera leaves extracts on the
Kim HK, Park HR, Lee JS, Chung TS, Chung HY, Chung lipopolysaccharide-activated Raw 264.7 cells,
J (2007) Down-regulation of iNOS and TNF-alpha pp 97–104. In: Green and smart technology with sen-
expression by kaempferol via NF-kappaB inactivation sor applications. Springer, Berlin/Heidelberg
in aged rat gingival tissues. Biogerontology La Cour B, Molgaard P, Yi Z (1995) Traditional Chinese
8(4):399–408 medicine in treatment of hyperlipidaemia.
Kim KH, Chang SW, Ryu SY, Choi SU, Lee KR (2009) J Ethnopharmacol 46(2):12–125
Phytochemical constituents of Nelumbo nucifera. Nat Lee KS, Lee KY (2011) Effect of lotus (Nelumbo nucifera)
Prod Sci 15(2):90–95 leaf extract on serum and liver lipid levels of rats fed a
Kim T, Kim HJ, Cho SK, Kang WY, Baek H, Jeon HY, high fat diet. J Korean Soc Food Sci Nutr
Kim BY, Kim D (2011) Nelumbo nucifera extracts as 40:1544–1547
104 Nelumbonaceae

Lee HK, Choi YM, Noh DO, Suh HJ (2005) Antioxidant nucifera on autoimmune disease of MRL/MpJ-lpr/lpr
effect of Korean traditional lotus liquor (Yunyupju). mice. Eur J Pharmacol 531(1–3):270–279
Int J Food Sci Technol 40(7):709–715 Liu CP, Kuo YC, Shen CC, Wu MH, Liao JF, Lin YL,
Lee HJ, Chen CC, Chou FP, Wu CH, Lai FS, Yang MY, Chen CF, Tsai WJ (2007) (S)-armepavine inhibits
Wang CJ (2010) Water extracts from Nelumbo nucifera human peripheral blood mononuclear cell activation by
leaf reduced plasma lipids and atherosclerosis in regulating Itk and PLCgamma activation in a PI-3K-
cholesterol-fed rabbits. J Food Biochem 34:779–795 dependent manner. J Leukoc Biol 81(5):1276–1286
Li GR, Li XG, Lu FH (1989) Effects of neferine on trans- Liu S, Wang B, Li XZ, Qi LF, Liang YZ (2009) Preparative
membrane potential in rabbit sinoatrial nodes and separation and purification of liensinine, isoliensinine
clusters of cultured myocardial cells from neonatal and neferine from seed embryo of Nelumbo nucifera
rats. Zhongguo Yao Li Xue Bao 10(4):328–331 (In Gaertn using high-speed counter-current chromatog-
Chinese) raphy. J Sep Sci 32(14):2476–2481
Li GR, Qian JQ, Lu FH (1990) Effects of neferine on heart Liu S, Li D, Huang B, Chen YX, Lu XC, Wang YW
electromechanical activity in anaesthetized cats. (2013) Inhibition of pancreatic lipase, α-glucosidase,
Zhongguo Yao Li Xue Bao 11(2):158–161 (In α-amylase, and hypolipidemic effects of the total fla-
Chinese) vonoids from Nelumbo nucifera leaves.
Li SS, Wu J, Chen LG, Du H, Xu YJ, Wang LJ, Zhang HJ, J Ethnopharmacol 149(1):263–269
Zheng XC, Wang LS (2014) Biogenesis of C-glycosyl Liu SH, Lu TH, Su CC, Lay IS, Lin HY, Fang KM, Ho TJ,
flavones and profiling of flavonoid glycosides in lotus Chen KL, Su YC, Chiang WC, Chen YW (2014)
(Nelumbo nucifera). PLoS One 9(10):e108860 Lotus leaf (Nelumbo nucifera) and its active
Liao CH, Lin JY (2012) Purification, partial characteriza- constituents prevent inflammatory responses in mac-
tion and anti-inflammatory characteristics of lotus rophages via JNK/NF-κB signaling pathway. Am
(Nelumbo nucifera Gaertn) plumule polysaccharides. J Chin Med 42(4):869–889
Food Chem 135(3):1818–1827 Luo XB, Chen B, Liu JJ, Yao SZ (2005) Simultaneous
Liao CH, Lin JY (2013a) Lotus (Nelumbo nucifera analysis of N-nornuciferine, O-nornuciferine, nucifer-
Gaertn) plumule polysaccharide ameliorates pancre- ine, and roemerine in leaves of Nelumbo nucifera
atic islets loss and serum lipid profiles in non-obese Gaertn by high-performance liquid chromatography–
diabetic mice. Food Chem Toxicol 58:416–422 photodiode array detection–electrospray mass spec-
Liao CH, Lin JY (2013b) Purified active lotus plumule trometry. Anal Chim Acta 538(1):129–133
(Nelumbo nucifera Gaertn) polysaccharides exert anti- Lv L, Jiang C, Li J, Zheng T (2012) Protective effects of
inflammatory activity through decreasing toll-like lotus (Nelumbo nucifera Gaertn) germ oil against car-
receptor-2 and -4 expressions using mouse primary bon tetrachloride-induced injury in mice and cultured
splenocytes. J Ethnopharmacol 147(1):164–173 PC-12 cells. Food Chem Toxicol 50(5):1447–1453
Lim SS, Jung YJ, Hyun SK, Lee YS, Choi JS (2006) Rat Ma WY, Lu YB, Hu RL, Chen JH, Zhang ZZ, Pan YJ
lens aldose reductase inhibitory constituents of Nelumbo (2010) Application of ionic liquids based microwave-
nucifera stamens. Phytother Res 20(10):825–830 assisted extraction of three alkaloids N-nornuciferine,
Lin HY, Kuo YH, Lin YL, Chiang W (2009a) Antioxidative O-nornuciferine, and nuciferine from lotus leaf.
effect and active components from leaves of lotus Talanta 80:1292–1297
(Nelumbo nucifera). J Agric Food Chem 57(15): Ma CJ, Wang JJ, Chu HM, Zhang XX, Wang ZH, Wang
6623–6629 HL, Li G (2014) Purification and characterization of
Lin MC, Kao SH, Chung PJ, Chan KC, Yang MY, Wang aporphine alkaloids from leaves of Nelumbo nucifera
CJ (2009b) Improvement for high fat diet-induced Gaertn and their effects on glucose consumption in
hepatic injuries and oxidative stress by flavonoid- 3T3-L1 adipocytes. Int J Mol Sci 15(3):3481–3494
enriched extract from Nelumbo nucifera leaf. J Agric Macko E, Douglas B, Weisbach JA, Waltz DT (1972)
Food Chem 57(13):5925–5932 Pharmacology of nuciferine and related aporphines.
Lin RJ, Wu MH, Ma YH, Chung LY, Chen CY, Yen CM Arch Int Pharmacodyn Ther 197:261–273
(2014) Anthelmintic activities of aporphine from Man JM, Cai JW, Cai CH, Xu B, Huai HY, Wei CX (2012)
Nelumbo nucifera Gaertn. cv. Rosa-plena against Comparison of physicochemical properties of starches
Hymenolepis nana. Int J Mol Sci 15(3):3624–3639 from seed and rhizome of lotus. Carbohydr Polym
Ling ZQ, Xie BJ, Yang EL (2005) Isolation, characteriza- 88(2):676–683
tion, and determination of antioxidative activity of Mani SS, Subramanian IP, Pillai SS, Muthusamy K (2010)
oligomeric procyanidins from the seedpod of Nelumbo Evaluation of hypoglycemic activity of inorganic con-
nucifera Gaertn. J Agric Food Chem 53:2441–2445 stituents in Nelumbo nucifera seeds on streptozotocin-
Liu CP, Tsai WJ, Lin YL, Liao JF, Chen CF, Kuo YC induced diabetes in rats. Biol Trace Elem Res
(2004) The extracts from Nelumbo nucifera suppress 138(1–3):226–237
cell cycle progression, cytokine genes expression, and Mathew M, Subramanian S (2014) In vitro screening for
cell proliferation in human peripheral blood mononu- anti-cholinesterase and antioxidant activity of metha-
clear cells. Life Sci 75(6):699–716 nolic extracts of ayurvedic medicinal plants used for
Liu CP, Tsai WJ, Shen CC, Lin YL, Liao JF, Chen CF, Kuo cognitive disorders. PLoS One 9(1):e86804
YC (2006) Inhibition of (S)-armepavine from Nelumbo
Nelumbo nucifera 105

Mazumder UK, Gupta M, Pramanik G, Mukhopadhyay Mukherjee D, Biswas A, Bhadra S, Pichairajan V,

RK, Sarkar S (1992) Antifertility activity of seed of Biswas T, Saha BP, Mukherjee PK (2010a) Exploring
Nelumbo nucifera in mice. Indian J Exp Biol the potential of Nelumbo nucifera rhizome on mem-
30(6):533–534 brane stabilization, mast cell protection, nitric oxide
Mon AA (2000) Use of Lotus (Nelumbo nucifera) for synthesis, and expression of costimulatory
nutrient retrieval from Pond Mud. M.Sc. thesis, Asian molecules. Immunopharmacol Immunotoxicol
Institute of Technology, Bangkok, Thailand 32(3):466–472
Mongkolrat S, Palanuvej C, Ruangrungsi N (2012) Mukherjee D, Khatua TN, Venkatesh P, Saha BP,
Quality assessment and liriodenine quantification of Mukherjee PK (2010b) Immunomodulatory potential
Nelumbo nucifera dried leaf in Thailand. Pharmacogn of rhizome and seed extracts of Nelumbo nucifera
J 4(32):24–28 Gaertn. J Ethnopharmacol 128(2):490–494
Morales MA, Bustamante SE, Brito G, Paz D, Cassels BK Mukherjee D, Kumar NS, Khatua T, Mukherjee PK
(1997) Cardiovascular effects of plant secondary (2010c) Rapid validated HPTLC method for estima-
metabolites norarmepavine, coclaurine and norcoclau- tion of betulinic acid in Nelumbo nucifera
rine. Phytother Res 12:103–109 (Nymphaeaceae) rhizome extract. Phytochem Anal
Mukherjee PK, Balasubramanian R, Saha K, Saha BP, Pal 21(6):556–560
M (1995a) Antibacterial efficiency of Nelumbo Mutreja A, Agarwal M, Kushwaha S, Chauhan A (2008)
nucifera (Nymphaeaceae) rhizomes extract. Indian Effect of Nelumbo nucifera seeds on the reproductive
Drugs 32(6):274–276 organs of female rats. Iranian J Reproduct Med
Mukherjee PK, Das J, Balasubramanian R, Kakali S, Pal 6(1):7–11
M, Saha BP (1995b) Antidiarrhoeal evaluation of Nadkarni KM, Nadkarni AK (1982) Indian Materia
Nelumbo nucifera rhizome extract. Indian J Pharmacol Medica with ayurvedic, Unani-Tibbi, siddha, allo-
27(4):262–264 pathic, homeopathic, naturopathic & home remedies,
Mukherjee PK, Pal SK, Saha K, Saha BP (1995c) vol 2, 2nd edn. Sangam Books, Bombay
Hypoglycaemic activity of Nelumbo nucifera Gaertn. Nakamura S, Nakashima S, Tanabe G, Oda Y, Yokota N,
(Fam. Nymphaeaceae) rhizome (methanolic extract) Fujimoto K, Matsumoto T, Sakuma R, Ohta T, Ogawa
in streptozotocin‐induced diabetic rats. Phytother Res K, Nishida S, Miki H, Matsuda H, Muraoka O,
9(7):522–524 Yoshikawa M (2013) Alkaloid constituents from
Mukherjee PK, Saha K, Giri S, Pal M, Saha B (1995d) flower buds and leaves of sacred lotus (Nelumbo
Antifungal screening of Nelumbo nucifera nucifera, Nymphaeaceae) with melanogenesis inhibi-
(Nymphaeaceae) rhizome extract. Indian J Microbiol tory activity in B16 melanoma cells. Bioorg Med
35:327–330 Chem 21(3):779–787
Mukherjee PK, Das J, Saha K, Giri SN, Pal M, Saha BP National Institute of Materia Medica (NIMM) (1999)
(1996a) Antipyretic activity of Nelumbo nucifera rhi- Selected medicinal plants in Vietnam, vol 2. Science
zome extract. Indian J Exp Biol 34(3):275–276 and Technology Publishing House, Hanoi, 460 pp
Mukherjee PK, Pal M, Saha K, Saha BP, Das J (1996b) Nguyen KH, Ta TN, Pham T, Nguyen QT, Pham H,
Diuretic activity of extract of the rhizomes of Nelumbo Mishra S, Nyomba BL (2012) Nuciferine stimulates
nucifera Gaertn. (Fam. Nymphaeaceae). Phytother insulin secretion from beta cells – An in vitro compari-
Res 10(5):424–425 son with glibenclamide. J Ethnopharmacol
Mukherjee PK, Saha K, Balasubramanian R, Pal M, Saha 142(2):488–495
BP (1996c) Studies on psychopharmacological effects Nishibe S, Tsukamoto H, Kinoshita H, Kitagawa S,
of Nelumbo nucifera Gaertn. rhizome extract. Sakushima A (1986) Alkaloids from embryo of the
J Ethnopharmacol 54(2–3):63–67 seed of Nelumbo nucifera. J Nat Prod 49(3):547–548
Mukherjee PK, Saha K, Das J, Pal M, Saha BP (1997a) Nishimura K, Horii S, Anahashi T, Sugimoto Y, Yamada
Studies on the anti-inflammatory activity of rhizomes J (2013) Synthesis and pharmacological activity of
of Nelumbo nucifera. Planta Med 63(4):367–369 alkaloids from embryo of lotus, Nelumbo nucifera.
Mukherjee PK, Saha K, Saha BP (1997b) Effect of Chem Pharm Bull 61(1):59–68
Nelumbo nucifera rhizome extract on blood sugar Niu CH, Wang Y, Liu JD, Wang JL, Xiao JH (2013)
level in rats. J Ethnopharmacol 58(3):207–213 Protective effects of neferine on amiodarone-induced
Mukherjee PK, Pal M, Pal SK, Shah K, Saha BP (1998) pulmonary fibrosis in mice. Eur J Pharmacol
Pharmacognostical profiles of rhizomes of Nelumbo 714(1–3):112–119
nucifera Gaertn. Anc Sci Life 17(4):273–279 Ochse JJ, Bakhuizen van den Brink RC (1980) Vegetables
Mukherjee PK, Kumar V, Houghton PJ (2007) Screening of the Dutch Indies, 3rd edn. Ascher & Co.,
of Indian medicinal plants for acetylcholinesterase Amsterdam, 1016 pp
inhibitory activity. Phytother Res 21(12):1142–1145 Ogle BM, Dao HTA, Mulokozi G, Hambraeus L (2001)
Mukherjee PK, Mukherjee D, Maji AK, Rai S, Heinrich Micronutrient composition and nutritional importance
M (2009) The sacred lotus (Nelumbo nucifera) – phy- of gathered vegetables in Vietnam. Int J Food Sci Nutr
tochemical and therapeutic profile. J Pharm Pharmacol 52:485–499
106 Nelumbonaceae

Oh JH, Choi BJ, Chang MS, Park SK (2009) Nelumbo generation in A549 cells. Food Chem
nucifera semen extract improves memory in rats with 141(4):3598–3605
scopolamine-induced amnesia through the induction Poornima P, Weng CF, Padma VV (2014) Neferine, an
of choline acetyltransferase expression. Neurosci Lett alkaloid from lotus seed embryo, inhibits human lung
461(1):41–44 cancer cell growth by MAPK activation and cell cycle
Ohara K, Wakabayashi H, Taniguchi Y, Shindo K, Yajima arrest. Biofactors 40(1):121–131
H, Yoshida A (2013) Quercetin-3-O-glucuronide Qian JQ (2002) Cardiovascular pharmacological effects
induces ABCA1 expression by LXRα activation in of bisbenzylisoquinoline alkaloid derivatives. Acta
murine macrophages. Biochem Biophys Res Commun Pharmacol Sin 23(12):1086–1092
441(4):929–934 Rahman W, Ilyas M, Khan AW (1962) Flower pigments:
Ohkoshi E, Miyazaki H, Shindo K, Watanabe H, Yoshida Flavonoids from Nelumbo nucifera Gaerten.
A, Yajima H (2007) Constituents from the leaves of Naturwissenschaften 49(14):327
Nelumbo nucifera stimulate lipolysis in the white adi- Rai S, Wahile A, Mukherjee K, Saha BP, Mukherjee PK
pose tissue of mice. Planta Med 73(12):1255–1259 (2006) Antioxidant activity of Nelumbo nucifera (sacred
Onishi E, Yamada K, Yamada T, Kaji K, Inoue H, Seyama lotus) seeds. J Ethonopharmacol 104(3):322–327
Y, Yamashita S (1984) Comparative effects of crude Sakuljaitrong S, Buddhakala N, Chomko S, Talubmook C
drugs on serum lipids. Chem Pharm Bull (2013) Effects of flower extract from lotus (Nelumbo
32(2):646–650 nucifera) on hypoglycemic and hypolipidemic in
Ono Y, Hattori E, Fukaya Y, Imai S, Ohizumi Y (2006) streptozotocin-induced diabetic rats. J Sci Eng Res
Anti-obesity effect of Nelumbo nucifera leaves 4(7):1441–1446
extract in mice and rats. J Ethnopharmacol Santhoshkumar T, Rahuman AA, Rajakumar G,
106(2):238–244 Marimuthu S, Bagavan A, Jayaseelan C, Zahir AA,
Ou M (1989) Chinese–English manual of common-used Elango G, Kamaraj C (2011) Synthesis of silver
prescriptions in traditional Chinese medicine. Joint nanoparticles using Nelumbo nucifera leaf extract and
Publishing Co, Ltd., Hong Kong, 753 pp its larvicidal activity against malaria and filariasis vec-
Pan PC, Chou YL, Sun TC, Kao YS (1962) Studies on the tors. Parasitol Res 108(3):693–702
alkaloids of embryo of loti Nelumbo nucifera. II Santhoshkumar T, Rahuman AA, Bagavan A, Kirthi AV,
Structure of liensinine. Sci China Math 5(3):321–336 Marimuthu S, Jayaseelan C, Kamaraj C, Zahir AA,
(In Chinese) Elango G, Rajakumar G, Velayutham K (2012)
Pan Y, Cai BC, Wang KL, Wang SM, Zhou SY, Yu XC, Efficacy of adulticidal and larvicidal properties of
Xu B, Chen L (2009) Neferine enhances insulin sensi- botanical extracts against Haemaphysalis bispinosa,
tivity in insulin resistant rats. J Ethnopharmacol Hippobosca maculata, and Anopheles subpictus.
124(1):98–102 Parasitol Res 111(4):1833–1840
Park YS, Towantakavanit K, Kowalska T, Jung ST, Ham Shad MA, Nawaz H, Hussain M, Yousuf B (2011)
KS, Heo BG, Cho JY, Yun JG, Kim HJ, Gorinstein S Proximate composition and functional properties of
(2009) Bioactive compounds and antioxidant and anti- rhizomes of lotus (Nelumbo nucifera) from Punjab,
proliferative activities of Korean white lotus cultivars. Pakistan. Pak J Bot 43(2):895–904
J Med Food 12(5):1057–1064 Shen-Miller J, Lindner P, Xie Y, Villa S, Wooding K,
Peng ZY, Zhang SD, Liu S, He BM (2011) Protective Clarke SG, Loo RRO, Loo JA (2013) Thermal-stable
effect of neferine on endothelial cell nitric oxide pro- proteins of fruit of long-living sacred lotus Nelumbo
duction induced by lysophosphatidylcholine: the role nucifera Gaertn var. China Antique. Trop Plant Biol
of the DDAH-ADMA pathway. Can J Physiol 6(2–3):69–84
Pharmacol 89(4):289–294 Shim SY, Choi JS, Byun DS (2009) Kaempferol isolated
Phillips R, Rix M (1993) Vegetables. Pan Books, London, from Nelumbo nucifera stamens negatively regulates
270pp FcepsilonRI expression in human basophilic KU812F
Phonkot N, Wangsomnuk PP, Aromdee C (2008) cells. J Microbiol Biotechnol 19(2):155–160
Antioxidant activity and DNA fingerprint of four vari- Shoji N, Umeyama A, Saito N, Iuchi A, Takemoto T,
eties of lotus stamens (Nelumbo nucifera Gaertn.). Kajiwara A, Ohizumi Y (1987) Asimilobine and lirini-
Songklanakarin J Sci Technol 30(1):55–58 dine, serotoninergic receptor antagonists, from
Phonkot N, Wangsomnuk PP, Aromdee C (2010) High- Nelumbo nucifera. J Nat Prod 50(4):773–774
performance liquid chromatographic determination of Shrestha MK, Knud-Hansen CF (1994) Increasing
β-carotene content in four varieties of lotus stamens. attached microorganism biomass as a management
Songklanakarin J Sci Technol 32(4):373–377 strategy for Nile tilapia (Oreochromis niloticus) pro-
Poornima P, Quency RS, Padma VV (2013a) Neferine duction. Aquacult Eng 13:101–108
induces reactive oxygen species mediated intrinsic Siegner R, Heuser S, Holtzmann U, Söhle J, Schepky A,
pathway of apoptosis in HepG2 cells. Food Chem Raschke T, Stäb F, Wenck H, Winnefeld M (2010)
136(2):659–667 Lotus leaf extract and l-carnitine influence different
Poornima P, Weng CF, Padma VV (2013b) Neferine from processes during the adipocyte life cycle. Nutr Metab
Nelumbo nucifera induces autophagy through the inhi- 7:66–75
bition of PI3K/Akt/mTOR pathway and ROS hyper
Nelumbo nucifera 107

Sinha S, Mukherjee PK, Mukherjee K, Pal M, Mandal SC, Nelumbo nucifera Gaertn. and Cyperus alternifolius
Saha BP (2000) Evaluation of antipyretic potential of L. J Environ Manage 137:54–60
Nelumbo nucifera stalk extract. Phytother Res Tian N, Liu Z, Huang J, Luo G, Liu S, Liu X (2007)
14(4):272–274 Isolation and preparation of flavonoids from the leaves
Sittiwet C (2009) Antimicrobial activity of essential oil of Nelumbo nucifera Gaertn by preparative reversed-
from Nelumbo nucifera Gaertn. Pollen. Int J Pharmacol phase high performance liquid chromatography. Se Pu
5(1):98–100 25(1):88–92 (In Chinese)
Smith RV, Sood SP (1971) In vitro metabolism of cer- Tomita M, Furukawa H (1962) On the alkaloids of
tain nornuciferine derivatives. J Pharm Sci 60(11): Nelumbo nucifera Gaertn. V. Alkaloids of Ohga-Hasu.
1654–1658 Yakugaku Zasshi 82(10):1458–1460 (In Japanese)
Sohn DH, Kim YC, Oh SH, Park EJ, Li X, Lee BH (2003) Tomita M, Watanabe Y, Furukawa H (1961a) On the alka-
Hepatoprotective and free radical scavenging effects of loids of Nelumbo nucifera Gaertn. II. Structure of nor-
Nelumbo nucifera. Phytomedicine 10(2–3):165–169 nuciferine. Yakugaku Zasshi 81(7):942–947 (In
Sridhar KR, Bhat R (2007) Lotus-a potential nutraceutical Japanese)
source. J Agric Technol 3(1):143–155 Tomita M, Watanabe Y, Furukawa H (1961b) On the alka-
Subashini R, Ponmurugan P, Rajadurai M (2012) Nelumbo loids of Nelumbo nucifera Gaertn. IV. Isolation of dl-
nucifera leaf extract ameliorates lipidperoxides and armepavine. Yakugaku Zasshi 81(11):1644–1647 (In
antioxidants in isoproterenolinduced myocardial Japanese)
infarction: biochemical and histological examination. Tomita M, Watanabe Y, Tomita M, Furukawa H (1961c)
J Pharm Res 5(7):3878–3882 On the alkaloids of Nelumbo nucifera Gaertn.
Sugimoto Y, Furutani S, Itoh A, Tanahashi T, Nakajima H, I. Yakugaku Zasshi 81(4):469–473 (In Japanese)
Oshiro H, Sun S, Yamada J (2008) Effects of extracts Tomita M, Furukawa H, Yang TH, Lin TJ (1964) Studies
and neferine from the embryo of Nelumbo nucifera on the alkaloids of loti embryo. I. Structure of isolien-
seeds on the central nervous system. Phytomedicine sinine. Tetrahedron Lett 5(37):2637–2642
15(12):1117–1124 Tomita M, Furukawa H, Yang TH, Lin TJ (1965) On the
Sugimoto Y, Furutani S, Nishimura K, Itoh A, Tanahashi alkaloids of Nelumbo nucifera Gaertn. VIII. Studies
T, Nakajima H, Oshiro H, Sun S, Yamada J (2010) on the alkaloids of loti embryo. (1). Structure of isoli-
Antidepressant-like effects of neferine in the forced ensinine, a new biscoclaurine type alkaloid. Chem
swimming test involve the serotonin1A (5-HT1A) Pharm Bull 13(1):39–43
receptor in mice. Eur J Pharmacol 634(1–3):62–67 Tomita M, Kunitomo J, Miyoshi M, Maruyama T, Yuge E
Sung JH, Sung JS, Shin HS (2011) Cytoprotective effects (1971) Studies on the Alkaloids of Nelumbo nucifera
of lotus (Nelumbo nucifera Gaertner) seed extracts on Gaertn. XV: synthesis of dl-Nornuciferine (dl-1-
oxidative damaged mouse embryonic fibroblast cell. Methoxy-2-hydroxyaporphine). Yakugaku Zasshi
Food Sci Biotechnol 20(6):1533–1537 91(8):896–900 (In Japanese)
Suthienkul O, Miyazaki O, Chulasiri M, Kositanont U, Tsuruta Y, Nagao K, Kai S, Tsuge K, Yoshimura T,
Oishi K (1993) Retroviral reverse transcriptase inhibi- Koganemaru K, Yanagita T (2011) Polyphenolic
tory activity in Thai herbs and spices: screening with extract of lotus root (edible rhizome of Nelumbo
Moloney murine leukemia viral enzyme. Southeast nucifera) alleviates hepatic steatosis in obese diabetic
Asian J Trop Med Public Health 24(4):751–755 db/db mice. Lipids Health Dis 10(1):202–230
Suzuki A, Kaneyama M, Shibanuma K, Takeda Y, Abe J, Tsuruta Y, Nagao K, Shirouchi B, Nomura S, Tsuge K,
Hizukuri S (1992) Characterization of lotus starch. Koganemaru K, Yanagita T (2012) Effects of lotus root
Cereal Chem 69(3):309–315 (the edible rhizome of Nelumbo nucifera) on the develop-
Talukder MJ, Nessa J (1998) Effect of Nelumbo nucifera ment of non-alcoholic fatty liver disease in obese diabetic
rhizome extract on the gastrointestinal tract of rat. db/db mice. Biosci Biotechnol Biochem 76(3):462–466
Bangladesh Med Res Counc Bull 24(1):6–9 U.S. Department of Agriculture, Agricultural Research
Tanaka Y, Nguyen VK (2007) Edible wild plants of Service (USDA-ARS) (2014) USDA national nutrient
Vietnam: the bountiful garden. Orchid Press, Bangkok, database for standard reference, release 26. Nutrient
175 pp data laboratoryhome page.
Tang CC, Lin WL, Lee YJ, Tang YC, Wang CJ (2014) ba/bhnrc/ndl
Polyphenol-rich extract of Nelumbo nucifera leaves Ushimaru T, Kanematsu S, Katayama M, Tsuji H (2001)
inhibits alcohol-induced steatohepatitis via reducing Antioxidative enzymes in seedlings of Nelumbo
hepatic lipid accumulation and anti-inflammation in nucifera germinated under water. Physiol Plant
C57BL/6 J mice. Food Funct 5(4):678–687 112(1):39–46
Tang W, Eisenbrand G (1992) Chinese drugs of plant ori- Ushimaru T, Hasegawa T, Amano T, Katayama M, Tanaka
gin. Springer, Berlin/Heidelberg/New York/London/ S, Tsuji H (2003) Chloroplasts in seeds and dark-grown
Paris/Tokyo/Hong Kong, 1065 pp seedlings of lotus. J Plant Physiol 160(3):321–324
Thongtha S, Teamkao P, Boonapatcharoen N, Tripetchkul Velusami CC, Agarwal A, Mookambeswaran V (2013)
S, Techkarnjararuk S, Thiravetyan P (2014) Effect of Nelumbo nucifera petal extracts on lipase,
Phosphorus removal from domestic wastewater by adipogenesis, adipolysis, and central receptors of obe-
108 Nelumbonaceae

sity. Evid Based Comp Alternat Med 2013. Article ID quality evaluation of Receptaculum Nelumbinis by
145925 RP-HPLC coupled with hierarchical clustering analy-
Wan JD, Zhao LB, Xu C, Zhang SH, Zhang ZX, Zeng C, sis. Int J Mol Sci 14(1):1999–2010
Chang MJ, Xiao JH, Wang JL (2011) Effects of nefer- Xiao JH, Zhang JH, Chen HL, Feng XL, Wang JL (2005)
ine on the pharmacokinetics of amiodarone in rats. Inhibitory effects of isoliensinine on bleomycin-
Biomed Chromatogr 25(8):858–866 induced pulmonary fibrosis in mice. Planta Med
Wang JL, Hu XM, Yin WH, Cai HS (1991) Studies on 71(3):225–230
alkaloids of Plumula Nelumbinis. Zhongguo Zhong Xiao JH, Zhang YL, Feng XL, Wang J, Qian JQ (2006)
Yao Za Zhi (J Chin Mat Med) 16(11):673–675, 703. Effects of isoliensinine on angiotensin II-induced pro-
(In Chinese) liferation of porcine coronary arterial smooth muscle
Wang JL, Nong Y, Jing MX (1992) Effects of liensinine cells. J Asian Nat Prod Res 8(3):209–216
on haemodynamics in rats and the physiologic proper- Xiao H, Chen AH, Ji AM, Li ZL, Song XD (2007)
ties of isolated rabbit atria. Yao Xue Xue Bao Isolation and identification of pronuciferine monomer.
27(12):881–885 (In Chinese) Zhong Yao Cai 30(3):289–291 (In Chinese)
Wang JL, Nong Y, Xia GJ, Yao WX, Jiang MX (1993) Xiao JS, Xie BJ, Cao YP, Wu H, Sun ZD, Xiao D (2012)
Effects of linesinine on slow action potentials in myo- Characterization of oligomeric procyanidins and
cardium and slow inward current in canine cardiac identification of quercetin glucuronide from lotus
Purkinje fibers. Yao Xue Xue Bao 28(11):812–816 (In (Nelumbo nucifera Gaertn.) seedpod. J Agric Food
Chinese) Chem 60(11):2825–2829
Wang YX, Liu B, Shi RB (2008) HPLC determination of Xie B, Wan J, Wang WQ, Shi CY, Hou XL, Fang JG
2-hydroxy-1-methoxyaporphine, pronuciferine, nucif- (2011) Nelumbo nucifera alkaloid inhibits 3 T3-L1
erine and roemerine in Nelumbo nucifera and its alka- preadipocyte differentiation and improves high-fat
loid fraction. Zhongguo Zhong Yao Za Zhi diet-induced obesity and body fat accumulation in
33(14):1713–1716 (In Chinese) rats. J Med Plants Res 5(10):2021–2028
Wang X, Liu JH, Geng YL, Wang DJ, Dong HJ, Zhang Xu X, Dong J, Mu XF, Sun LP (2011a) Supercritical CO2
TY (2010) Preparative separation of alkaloids from extraction of oil, carotenoids, squalene and sterols
Nelumbo nucifera Gaertn by pH-zone-refining counter- from lotus (Nelumbo nucifera Gaertn) bee pollen.
current chromatography. J Sep Sci 33(4–5):539–544 Food Bioprod Process Trans Inst Chem Eng Part C
Wang HM, Yang WL, Yang SC, Chen CY (2011) Chemical 89(1):47–52
constituents from the leaves of Nelumbo nucifera Gaertn. Xu X, Sun CR, Dai XJ, Hu RL, Pan YJ, Yang ZF (2011b)
cv. Rosa-plena. Chem Nat Comp 47(2):316–318 LC/MS guided isolation of alkaloids from lotus leaves
Wassel G, Saeed A, Ibrahim N, El-Eraqy W (1996) by pH-zone-refining counter-current chromatography.
Flavonoids of Nelumbo nucifera Gaertn and biological Molecules 3:2551–2560
evaluation. Egypt J Pharm Sci 37(1–6):585–595 Xu SH, Sun Y, Jing F, Duan WJ, Du JH, Wang X (2011c)
Weng TC, Shen CC, Chiu YT, Lin YL, Kuo CD, Huang Separation and purification of flavones from Nelumbo
YT (2009) Inhibitory effects of armepavine against nucifera Gaertn. by silica gel chromatography and
hepatic fibrosis in rats. J Biomed Sci 16:78 high-speed counter-current chromatography. Se Pu
Wongwattanasathien O, Kangsadalampai K, Tongyonk L 29(12):1244–1248 (In Chinese)
(2010) Antimutagenicity of some flowers grown in Yan SL, Wang QZ, Peng GH (2009) Determination of cat-
Thailand. Food Chem Toxicol 48(4):1045–1051 echin in lotus rhizomes by high-performance liquid
Wu MJ, Wang L, Weng CY, Yen JH (2003) Antioxidant chromatography. Int J Food Sci Nutr 60(5):432–438
activity of methanol extract of the lotus leaf Yang TH, Chen CM (1970a) Isolation of methylcorypall-
(Nelumbo nucifera Gertn.). Am J Chin Med ine from embryo loti. J Chin Chem Soc (Taipei)
31(5):687–698 17(1):54–56
Wu SH, Sun CR, Cao XJ, Zhou H, Hong Z, Pan YJ (2004) Yang TH, Chen CM (1970b) On the alkaloids of Nelumbo
Preparative counter-current chromatography isolation nucifera Gaertn. Studies on the alkaloid of loti embryo.
of liensinine and its analogues from embryo of the J Chin Chem Soc (Taipei) 17(4):113–120
seed of Nelumbo nucifera GAERTN. using upright Yang TH, Chen CM, Lu CS, Liao CL (1972) Studies on
coil planet centrifuge with four multilayer coils con- the alkaloids of lotus receptacle. J Chin Chem Soc
nected in series. J Chromatogr A 1041(1–2):153–162 19(3):143–147 (In Chinese)
Wu CH, Yang MY, Chan KC, Chung PJ, Ou TT, Wang CJ Yang DM, Wang QS, Ke LQ, Jiang JM, Ying TJ (2007)
(2010) Improvement in high-fat diet-induced obesity Antioxidant activities of various extracts of lotus
and body fat accumulation by a Nelumbo nucifera leaf (Nelumbo nuficera Gaertn) rhizome. Asia Pac J Clin
flavonoid-rich extract in mice. J Agric Food Chem Nutr 16(Suppl 1):158–163
58(11):7075–7081 Yang WM, Shim KJ, Choi MJ, Park SY, Choi BJ, Chang
Wu YB, Zheng LJ, Wu JG, Chen TQ, Yi J, Wu JZ (2012) MS, Park SK (2008) Novel effects of Nelumbo
Antioxidant activities of extract and fractions from nucifera rhizome extract on memory and neurogenesis
receptaculum nelumbinis and related flavonol glyco- in the dentate gyrus of the rat hippocampus. Neurosci
sides. Int J Mol Sci 13(6):7163–7173 Lett 443(2):104–107
Wu YB, Zheng LJ, Yi J, Wu JG, Chen TQ, Wu JZ (2013) Yang RZ, Wei XL, Gao FF, Wang LS, Zhang HJ, Xu YJ,
Quantitative and chemical fingerprint analysis for the Li CH, Ge YX, Zhang JJ, Zhang J (2009) Simultaneous
Nelumbo nucifera 109

analysis of anthocyanins and flavonols in petals of Yu LS, Shen Q, Zhou Q, Jiang HD, Bi HC, Huang M,
lotus (Nelumbo) cultivars by high-performance liquid Zhou H, Zeng S (2013b) In vitro characterization of
chromatography-photodiode array detection/electro- ABC transporters involved in the absorption and distri-
spray ionization mass spectrometry. J Chromatogr A bution of liensinine and its analogs. J Ethnopharmacol
1216:106–112 150(2):485–491
Yang MY, Chang YC, Chan KC, Lee YJ, Wang CJ (2011) Zhang XY, Liu ZJ, Xu B, Sun ZL, Gong YQ, Shao CS
Flavonoid-enriched extracts from Nelumbo nucifera (2012) Neferine, an alkaloid ingredient in lotus seed
leaves inhibits proliferation of breast cancer in vitro embryo, inhibits proliferation of human osteosarcoma
and in vivo. Eur J Integr Med 3(3):e153–e163 cells by promoting p38 MAPK-mediated p21 stabili-
Yang ZD, Zhang X, Du J, Ma ZJ, Guo F, Li S, Yao XJ zation. Eur J Pharmacol 677(1–3):47–54
(2012) An aporphine alkaloid from Nelumbo nucifera Zhang Y, Wang Y, Zheng BD, Lu X, Zhuang WJ (2013a)
as an acetylcholinesterase inhibitor and the primary The in vitro effects of retrograded starch (resistant
investigation for structure–activity correlations. Nat starch type 3) from lotus seed starch on the proliferation
Prod Res 26(5):387–392 of Bifidobacterium adolescentis. Food Funct
Ye LH, He XX, Kong LT, Liao YH, Pan RL, Xiao BX, Liu 4(11):1609–1616
XM, Chang Q (2014a) Identification and characteriza- Zhang YJ, Hao XY, Liang ZS, Ke WD, Guo HB (2013b)
tion of potent CYP2D6 inhibitors in lotus leaves. Efficient isolation of high-quality RNA from lotus
J Ethnopharmacol 153(1):190–196 Nelumbo nucifera ssp nucifera tissues. Genet Mol Res
Ye LH, He XX, Yan MZ, Chang Q (2014b) Identification 12(1):223–229
of in vivo components in rats after oral administration Zhang Y, Zeng HL, Wang Y, Zeng SX, Zheng BD (2014)
of lotus leaf flavonoids using ultra fast liquid chroma- Structural characteristics and crystalline properties of
tography with tandem mass spectrometry. Anal lotus seed resistant starch and its prebiotic effects.
Methods 6:6088–6094 Food Chem 155:311–318
Yen GC, Duh PD, Su HJ, Yeha CT, Wu CH (2006) Zhao X, Shen J, Chang KJ, Kim SH (2013a) Analysis of
Scavenging effects of lotus seed extracts on reactive Fatty acids and phytosterols in ethanol extracts of
nitrogen species. Food Chem 94(4):596–602 Nelumbo nucifera seeds and rhizomes by GC-MS. J
Yoo DY, Kim W, Yoo KY, Lee CH, Choi JH, Kang IJ, Agric Food Chem 61(28):6841–6847
Yoon YS, Kim DW, Won MH, Hwang IK (2011) Zhao XL, Wang ZM, Ma XJ, Jing WG, Liu A (2013b)
Effects of Nelumbo nucifera rhizome extract on cell Chemical constituents from leaves of Nelumbo
proliferation and neuroblast differentiation in the hip- nucifera. Zhongguo Zhong Yao Za Zhi 38(5):703–708
pocampal dentate gyrus in a scopolamine-induced (In Chinese)
amnesia animal model. Phytother Res 25(6):809–815 Zhao X, Shen J, Chang KJ, Kim SH (2014) Comparative
Yoon JS, Kim HM, Yadunandam AK, Kim NH, Jung HA, analysis of antioxidant activity and functional compo-
Choi JS, Kim CY, Kim GD (2013) Neferine isolated nents of the ethanol extract of lotus (Nelumbo nucifera)
from Nelumbo nucifera enhances anti-cancer activities from various growing regions. J Agric Food Chem
in Hep3B cells: molecular mechanisms of cell cycle 62(26):6227–6235
arrest, ER stress induced apoptosis and anti-angiogenic Zheng ZJ, Wang ML, Wang DJ, Duan WJ, Wang X,
response. Phytomedicine 20(11):1013–1022 Zheng CC (2010) Preparative separation of alkaloids
You JS, Lee YJ, Kim KS, Kim SH, Chang KJ (2014a) Anti‐ from Nelumbo nucifera leaves by conventional and
obesity and hypolipidaemic effects of Nelumbo nucifera pH-zone-refining counter-current chromatography.
seed ethanol extract in human pre‐adipocytes and rats J Chromatogr B Analyt Technol Biomed Life Sci
fed a high‐fat diet. J Sci Food Agric 94(3):568–575 878(19):1647–1651
You JS, Lee YJ, Kim KS, Kim SH, Chang KJ (2014b) Zhong G, Chen ZD, WeYM (2007) Physicochemical
Ethanol extract of lotus (Nelumbo nucifera) root properties of lotus (Nelumbo nucifera Gaertn.) and
exhibits an anti-adipogenic effect in human pre- kudzu (Pueraria hirsuta Matsum.) starches. Int J Food
adipocytes and anti-obesity and anti-oxidant effects in Sci Technol 42(12):1449–1455
rats fed a high-fat diet. Nutr Res 34(3):258–267 Zhou MG, Jiang M, Ying XH, Cui QX, Han YQ, Hou YY,
Youn UJ, Lee JH, Lee YJ, Nam JW, Bae H, Seo EK (2010) Gao J, Bai G, Luo GA (2013a) Identification and com-
Regulation of the 5‐HT3A receptor‐mediated current parison of anti-inflammatory ingredients from differ-
by alkyl 4‐hydroxybenzoates isolated from the seeds of ent organs of Lotus Nelumbo by UPLC/Q-TOF and
Nelumbo nucifera. Chem Biodivers 7(9):2296–2302 PCA coupled with a NF-κB reporter gene assay. PLoS
Yu J, Hu WS (1997) Effects of neferine on platelet aggre- One 8(11):1
gation in rabbits. Yaoxue Xuebao 32:1–4 (In Chinese) Zhou YJ, Xiang JZ, Yuan H, Liu H, Tang Q, Hao HZ, Yin
Yu HG, Cheng LB, Yin JJ, Yan SJ, Liu KJ, Zhang FM, Xu Z, Wang J, Ming ZY (2013b) Neferine exerts its anti-
B, Li LJ (2013a) Structure and physicochemical prop- thrombotic effect by inhibiting platelet aggregation
erties of starches in lotus (Nelumbo nucifera Gaertn.) and promoting dissociation of platelet aggregates.
rhizome. Food Sci Nutr 1(4):273–283 Thromb Res 132(2):202–210
Mirabilis expansa

Scientific Name Spanish: Arracacha De Toro, Arricón, Camotillo,

Chagos, Cushpe, Mauka, , Pega-Pega, Shallca
Mirabilis expansa (Ruiz & Pav.) Standl. Yuca, Yuca De La Jalca, Yuca Inca

Allionia expansa (Ruiz & Pav.) Kuntze,
Calyxhymenia expansa Ruiz & Pav., Oxybaphus Mauka is indigenous to the Andes in South
expansa (Ruiz & Pav.) Vahl America, distributed naturally in the area of La Paz
(Bolivia), to the North of Quito (Ecuador) and in
Cajamarca (Peru); however, it was also found in
Family Venezuela and Chile (Popenoe et al. 1989; Bermejo
and León 1994; Flores 2003). Its wild ancestors are
Nyctaginaceae found in Peru, Bolivia, Ecuador and Colombia.

Common/English Names Agroecology

Chago, Mauka Mauka is found in the cold, windy altitudes above

2700–3500 m in the Andes where other crops
like potato cannot survive the harsh ecological
Vernacular Names conditions (Popenoe et al. 1989; Bermejo and
León 1994). The area is characterised by mean
Bolivia: Mauka annual temperature of 13 °C and with maxima of
Ecuador: Miso, Pega-Pega, Taso 25 °C and minima of 5 °C and an annual precipi-
Peru: Chago, Arricón, Yuca, Inca, Cushipe, tation of 680 mm and intensive sunshine.
Chaco However, in Cajamarca, it thrives at 600–
1000 mm per year. It thrives well on fertile,

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DOI 10.1007/978-3-319-26062-4_6
Mirabilis expansa 111

moist, deep, loose alluvial soils with abundant

organic matter (Flores 2003).

Edible Plant Parts and Uses

The lower stem parts, swollen roots cooked and

leaves are edible (Popenoe et al. 1989; Facciola
1990; Bermejo and León 1994). The stem and
swollen roots are usually boiled or fried as vege-
tables. Freshly harvested, they contain an astrin-
gent principle which can be removed by drying
them in the sun. Traditionally, the dried and sun-
Plate 1 Mauka tuberous root (Frank Van Keirsbilck)
sweetened tubers are chopped, boiled and mixed
with honey or brown sugar and toasted grain in
Bolivia. The combined ingredients afford a
hearty meal, and the cooking water makes an
especially flavourful drink. In Ecuador, mauka is
processed in two ways, salty and sweet (de sal or
de dulce). For salty mauka, the tubers are cleaned,
cooked and peeled and then eaten immediately or
used as an ingredient of soups and stews. To
make sweet mauka, the stems and roots are lay-
ered with barley or mauka stems in a pit dug in
the ground for about several weeks to a month,
by which time the starches have largely hydro-
lysed to sugars. Both salty and sweet forms are
commonly mixed with syrup or molasses and
eaten with tomatoes and fish (particularly sar- Plate 2 Mauka foliage (Frank Van Keirsbilck)
dines or tuna). The leaves are also eaten as a
cooked leafy vegetable or used raw in salads and
chilli sauces. with reddish edges. The inflorescences are termi-
nal racemes, 3–6 cm long covered with viscid
hairs. Flowers apetalous, calyx tubular with
Botany 5-cleft purple, white or white-purple mottled
perianth; stamens 5–6 as long as perianth;
Mirabilis expansa is a low, compact, herbaceous filaments are connate at the base; style 1 with a
perennial, growing to 1 m high. The swollen capitate stigma. Fruit is hard, capsule like and
stems below ground are white, salmon coloured indehiscent. Seeds are small and dark brown.
or yellow. They are commonly smooth and fleshy,
about 5 cm in diameter and 50 cm in length.
Young tuberous roots are yellow and older ones Nutritive/Medicinal Properties
white (Plate 1). The aerial portion is made up of
a mass of foliage formed from the basal shoots. Chemical composition of mauka roots (per 100-g
The aerial stems are cylindrical, with short dis- dry matter) was reported as: energy 427 kcal,
tinct internodes, pale green or with reddish mark- moisture 61.94 %, fibre 4.83 %, protein 7.41 %,
ings with opposite ovate or cordate, pubescent, ash 4.49 %, starch 67.71 %, carbohydrates
petioled leaves (Plate 2), 8 cm by 3 cm wide often 80.46 %, Ca 0.61 %, P 0.09 %, Mg 0.09 %, Na
112 Nyctaginaceae

0.03 %, K 1.27 %, Cu 6 ppm, Fe 50 ppm, Mn Ribosome-inactivating protein (RIP) isolated

7 ppm and Zn 62 ppm (Klásková and Fernandéz from M. expansa showed substantial inhibitory
2011). activity against fungal growth (Vepachedu et al.
Bermejo and León (1994) reported that 2003). RIP from M. expansa was targeted to the
Bolivian maukas contained 7 % protein content, surface of fungal cells and transferred into the
2 760-mg Ca and 590-mg P (in dry matter) in the cells where they caused ribosome depurination
underground parts and a 17 % protein content in and subsequent fungal mortality. Two novel type
the foliage. They also reported that analysis of I ribosome-inactivating proteins (RIPs) were
three cultivars of chago from Cajamarca in Peru found in the storage roots of Mirabilis expansa
showed 4–5 % protein, 157 46-mg Ca and 117- and named ME1 and ME2 (Vivanco et al. 1999).
mg P per 100-g edible portion. Chago was found The two proteins were found to be similar in size
to be low in Fe and Na. (27 and 27.5 kD), and their isoelectric points
Popenoe et al. (1989) reported both the were determined to be greater than pH 10.0. ME2
swollen stems and the roots of mauka to be rich showed high similarity to the Mirabilis jalapa
in carbohydrates (87 % per dry weight basis) antiviral protein, a type I RIP. ME1 and ME2
with 7 % protein (an appreciable amount for a were active against several fungi, including
root crop) and little fibre. Based on an evalua- Pythium irregulare, Fusarium oxysporum solani,
tion of three separate ecotypes, mauka was Alternaria solani,Trichoderma reesei and
found to be richer than the other Andean tubers Trichoderma harzianum, and an additive antifun-
in Ca, K and P. The leaves were found to con- gal effect of ME1 and ME2 was observed.
tain about 17 % protein with higher level of Antibacterial activity of both ME1 and ME2 was
digestibility than that of the other forages observed against Pseudomonas syringae,
grown in the upland Andes. Agrobacterium tumefaciens, Agrobacterium
ME1, a type 1 ribosome-inactivating proteins radiobacter and others.
(RIP), was cloned and sequenced from storage
roots of Mirabilis expansa (Vepachedu et al.
2003, 2005). The full-length cDNA sequence of Other Uses
ME1 had 1129 nucleotides with an open reading
frame of 951 nucleotides representing 317 amino The whole plant is used as animal feed especially
acids. ME1 showed very close similarities to for guinea pigs and pigs (Popenoe et al. 1989).
MAP and MAP-4 from Mirabilis jalapa. ME1 Animals consume it fresh or dried. In feeding
was produced in large quantities in Mauka stor- pigs and guinea pigs, the raw tubers, leaves and
age roots. RIPs are toxic proteins synthesised by stems are often mixed with corn and weedy veg-
many plants and some bacteria, which specifi- etation, while cavies are fed the green foliage or
cally depurinate the 28S RNA and thus interrupt hay.
protein translation. RIPs hold broad interest
because of their potential use as plant defence
factors against pathogens. ME1 was shown to Comments
depurinate a variety of partially denatured nucleic
acids, randomly removing adenine residues from The plant is propagated from seeds or stem
single-stranded regions and, to a lesser extent, cuttings.
guanine residues from wobble base pairs in hair-
pin stems (Park et al. 2004). Exposure to ME1
in-vitro completely abolished the infectivity of Selected References
partially denatured RNA transcripts of the potato
spindle tuber viroid, suggesting that RIPs may Bermejo JEH, León J (eds) (1994) Neglected crops: 1492
target invading nucleic acids before they reach from a different perspective, Plant production and pro-
host ribosomes in-vivo. tection series no. 26. FAO, Rome, pp 205–209
Mirabilis expansa 113

Davidson A (2006) The Oxford companion to food, 2nd known plants of the Andes with promise for
edn. Oxford University Press, Oxford. 907 pp worldwide cultivation. National Academy Press,
Facciola S (1990) Cornucopia. A source book of edible Washington, DC, pp 37–46
plants. Kampong Publications, Vista. 677 pp The Plant List (2013) Version 1.1. Published on the
Flores H (2003) Andean root and tuber crops: under- Internet;
ground rainbows. Hort Sci 38(2):161–167 Vepachedu R, Bais HP, Vivanco JM (2003) Molecular
Klásková T, Fernandéz CE (2011) Mauka – a promise for characterization and post-transcriptional regulation of
fighting with the unbalanced nutrition of high Andean ME1, a type-I ribosome-inactivating protein from
regions. Agric Trop Subtrop 44(1):37–40 Mirabilis expansa. Planta 217(3):498–506
Park SW, Vepachedu R, Owens RA, Vivanco JM (2004) Vepachedu R, Park SW, Sharma N, Vivanvo JM (2005)
The N-glycosidase activity of the ribosome- Bacterial expression and enzymatic activity analysis
inactivating protein ME1 targets single-stranded of ME1, a ribosome-inactivating protein from
regions of nucleic acids independent of sequence or Mirabilis expansa. Protein Expr Purif 40:142–151
structural motifs. J Biol Chem 279(33): Vivanco JM, Savary BJ, Flores HE (1999) Characterization
34165–34174 of two novel type I ribosome-inactivating proteins
Popenoe H, King SR, Léon J, Kalinkowski LS, Vietmeyer from the storage roots of the Andean crop Mirabilis
ND, Dafforn M (1989) Lost crops of the Incas. Little expansa. Plant Physiol 119(4):1447–1456
Nymphaea odorata

Scientific Name Vernacular Names

Nymphaea odorata Aiton

French: Nymphéa Odorant, Nymphéa Tubéreux
German: Wohlriechende Seerose
Synonyms Swedish: Doftnäckros

Castalia minor (Sims) DC. ex Small, Nymphaea

odorata var. minor Sims, Nymphaea odorata var.
odorata, Nymphaea odorata f. odorata, Origin/Distribution
Nymphaea odorata subsp. odorata, Nymphaea
odorata var. rosea Pursh, Nymphaea odorata var. The species is native to North America (USA and
stenopetala Fernald, Nymphaea parkeriana Canada), Central America and the Caribbean
Lehm. (Grin 2014). The species occurs from Puerto
Rico to Alaska and from California to Quebec
(Kartesz 1999).

Nymphaeaceae Agroecology

Nymphaea odorata grows in shallow ponds,

Common/English Names lakes and their margins, ditches, swamps, slow
streams from sea level to 1700 m elevation. The
American White Water Lily, Beaver Root, species grows rooted in mucky or silty sediments
Fragrant Water Lily, Sweet-Scented Water Lily, in water up to 6–7 feet deep and can survive in
Tuberous Water Lily, White Water Lily both acid and alkaline waters. N. odorata plants

© Springer International Publishing Switzerland 2016 114

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_7
Nymphaea odorata 115

can be established from seeds in up to 90 cm (8) (Zhang et al. 2003). Compounds 3, 4 and 7
water and the seedlings grow as submerged showed marginal inhibitory effect against fatty
aquatics until they switch abruptly to the produc- acid synthase with IC50 values of 45, 50 and
tion of floating leaves (Richards and Cao 2012). 25 μg/mL, respectively.
Five plant growth inhibitory allelochemicals
from N. odorata were identified as gallic acid,
Edible Plant Parts and Uses myricitrin, myricetin, 1,2,3,4,6-pentagalloyl-D-
glucose and 2,3,4,6-tratragalloyl-D-glucose
Young leaves are used in soups and stews and (Elakovich et al. 1999).
tubers are also edible (Fernald et al. 1958; Tanaka The crude leaf extract of N. odorata exhibited
1976; Kunkel 1984; Facciola 1990). Flower buds weak larvicidal activity against the larvae of the
pickled or eaten as cooked vegetable. malaria vector, Anopheles gambiae, of 10 and
20 % (at 5 % w/v) and 20 and 30 % (at 10 % w/v)
both at 12 and 24 h incubation, respectively
Botany (Oladimeji et al. 2008). The crude leaf extract
and fractions were inactive against the bacterial
An aquatic perennial herbaceous plant with float- and fungal isolates tested.
ing leaves and branched creeping, fleshy rhi- Studies by Deoda et al. (2012) reported that
zomes. The rhizomes are attached by cluster of extracts of the leaf and bulbs exhibited better
adventitious roots arising below the leaf bases; antilithiatic activity than the stem extract of the
they bear circular leaf scars on the upper surface; Dolichos lablab bean extracts.
the rhizomes are brown externally, greyish-white
internally, spongy and are densely covered with
short black hairs. Leaves are circular (10–35 cm Traditional Medicinal Uses
across), cleft at the base, with entire, undulating
margin, shining green and waxy above, reddish The root has been reported to be alterative,
or purplish and more or less hairy on under sur- anodyne, antiseptic, astringent and demulcent
face, petiole slender and cylindrical attached to (Grieve 1971; Lust 1974; Mill 1985; Bown
centre of lower leaf surface (Plate 1). Flowers 1995). A tea made from the roots is used in the
solitary up to 25 cm across, on a grooved pedun- treatment of tuberculosis, chronic bronchial
cle, floating on water surface or raised, with 4 complaints, diarrhoea, dysentery, gastrointestinal
green sepals and 15–40 white or pinkish, lanceo- inflammation, gonorrhoea, vaginal discharge,
late petals with acute apices, 55–120 yellow sta- inflamed glands, mouth sores and to stop bleed-
mens (Plate 1). Fruit an ovoid berry-like capsule, ing (Bown 1995; Foster and Duke 1998). A poul-
1–2 cm across and containing many seeds tice made from the roots is used in the treatment
(2–3 mm long). of swellings, boils, tumours, inflamed skin and
vaginitis (Bown 1995; Foster and Duke 1998).
Moerman (1998) recorded the following
Nutritive/Medicinal Properties ethnomedicinal uses of the plant by American
natives: The Chippewa use pulverized roots for
The following compounds were identified from mouth sores; the Mimac use leaves as cold rem-
fractionation of the ethanol extract of N. odorata: edy, the juice of root is taken for coughs and
nymphaeoside A (1), icariside E(4) (2), kaempferol poultice of boiled roots applied to swellings. The
3-O-α-L-rhamnopyranoside (afzelin, 3), querce- Ojibwe use the root as cough medicine and for
tin 3-O-α-L-rhamnopyranoside (4), myricetin tuberculosis. The Okanagan-Colville place stems
3-O-α-L-rhamnopyranoside (myricitrin, 5), quer- directly onto tooth for toothache. The Penobscot
cetin 3-O-(6″-O-acetyl)-β-D-galactopyranoside apply leaves as poultice on limb swellings. The
(6), myricetin 3-O-β-D-galactopyranoside (7) and Potawatomi used poultice of pounded roots for
myricetin 3-O-(6″-O-acetyl)-β-D-galactopyranoside unspecified ailments.
116 Nymphaeaceae

Plate 1 White water lily flower

and leaves

Nymphaea odorata is an old herbal recipe and stem of Nymphaea odorata and Dolichos lablab
beans. Res J Pharm Biol Chem Sci 3(1):814–819
used in the treatment and/or management of
Elakovich S, Spence S, Yang J (1999) Chapter 4:
ocular, skin, gastrointestinal and urino-genital Phytochemical inhibitors from the Nymphaeceae:
ailments amongst many others (Oladimeji et al. Nymphaea odorata and Nuphar lutea. In: Cutler HG,
2008). Nymphaea odorata has been traditionally Cutler SJ (eds) Biologically active natural products.
CRC Press, Boca Raton, pp 49–56
deemed to be employed as antiseptic, astringent,
Facciola S (1990) Cornucopia. A source book of edible
demulcent, kidney stone, deobstruent, discutient plants. Kampong Publications, Vista. 677 pp
and vulnerary (Deoda et al. 2012). Roots are used Fernald ML, Kinsey AX, Rollins RC (1958) Wild plants
for coughs, mouth sores and for tuberculosis. of Eastern North America, 2nd edn. Harper & Row,
New York
Foster S, Duke JA (1998) A field guide to medicinal plants
in Eastern and Central N. America. Houghton Mifflin
Other Uses Co, Boston. 366 pp
Grieve M (1971) A modern herbal. Penguin. 2 vols. Dover
publications, New York, 919 pp
N. odorata is a common and popular ornamental
Kartesz JT (1999) A synonymized checklist and atlas with
for ponds. biological attributes for the vascular flora of the United
States, Canada, and Greenland. First edition. In:
Kartesz JT, Meacham CA (eds) Synthesis of the North
American Flora, Version 1.0. North Carolina Botanical
Comments Garden, Chapel Hill
Kunkel G (1984) Plants for human consumption. An
A declared aquatic or terrestrial noxious weed annotated checklist of the edible phanerogams and
and/or noxious-weed seed in some states in the ferns. Koeltz Scientific Books, Koenigstein
Lust JB (1974) The herb book. Bantam Books, New York.
USA. Dense infestations may also accelerate the
174 pp
natural siltation process in the shallow bodies of Mills SY (1985) The dictionary of modern herbalism.
water. Thorsons, Wellingborough
Moerman DE (1998) Native American ethnobotany.
Timber Press, Portland. 927 pp
Oladimeji HO, Ubulom PM, Akpabio EI, Etim IE, Nyong
Selected References E (2008) Larvicidal and anti-microbial potentials of
Nymphaea odorata. J Pharmacol Toxicol 3(5):
Bown D (1995) Encyclopaedia of herbs and their uses. 357–362
Dorling Kindersley, London. 424 pp Richards JH, Cao C (2012) Germination and early growth
Deoda RS, Pandya H, Patel M, Yadav KN, Kadam PV, of Nymphaea odorata at different water depths. Aquat
Patil MJ (2012) Antilithiatic activity of leaves, bulb Bot 98(1):12–19
Nymphaea odorata 117

Tanaka T (1976) Tanaka’s cyclopaedia of edible plants of Wiersema JH (1997) Nymphaea. In: Flora of North
the world. Keigaku Publishing, Tokyo. 924 pp America Editorial Committee (eds) 1993+. Flora of
The Plant List (2013) Version 1.1. Published on the Internet; North America North of Mexico. 19+ vols. New York and Oxford
USDA, ARS, National Genetic Resources Program (GRIN) Zhang ZZ, ElSohly HN, Li XC, Khan SI, Broedel SE Jr,
(2014) Germplasm Resources Information Network – Raulli RE, Cihlar RL, Burandt C, Walker LA (2003)
(GRIN) [Online Database]. National Germplasm Phenolic compounds from Nymphaea odorata. J Nat
Resources Laboratory, Beltsville, Maryland. http:// Prod 66:548–550
Nymphaea x rubra

Scientific Name (Hindu), Tharo Angangba (Manipur), Lal

Kamal, Raktakamal (Marathi), Li’ne Aluk
Nymphaea x rubra Roxb. ex Andrews (see ori- (Mising, Assam), Alagandha, Alipriya,
gin/distribution) Alohita, Kumuda, Raktakamal (Sanskrit) Atti
(Tamil), Ranga Podum (Tai-Khamyang,
Assam), Erra Kaluva, Thaamara (Telugu),
Synonyms Nilofar (Urdu)
Japanese: Hime-Suiren
Castalia rubra (Roxb. ex Andrews) Tratt. Portuguese: Ninfeia Vermelha
Spanish: Nenufar Rojo
Vietnamese: Bông Súng Ðỏ


Common/English Names The species is reported to be indigenous to the

Indian subcontinent, naturalised in tropical Asia
Red Water Lily, Red Indian Lily, Mini Water Lily, and cultivated in the Caribbean and elsewhere
Dwarf Lily, Red Lily of the Nile (USDA-ARS 2015). However, Conrad (1905) in
his monograph of the genus Nymphaea expressed
his doubts whether true Nymphaea rubra existed
Vernacular Names in India.
Recent studies by Dkhar et al. (2013) using
Burmese: Krani, Kyar-Ni molecular cloning and sequencing of ITS (inter-
French: Lotus Rouge, Nenuphar Rouge nal transcribed spacer) region confirmed
German: Rote Seerose Nymphaea rubra to be a hybrid produced through
India: Mokua, Ronga Bhet, Ronga Seluk, Seluk a cross between N. lotus and N. pubescens with
(Assamese), Lal Shapla (Bengali), Kokaa the latter representing the maternal parent. Based

© Springer International Publishing Switzerland 2016 118

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_8
Nymphaea x rubra 119

on these evidences and its inability to form seeds Botany

in nature, they proposed that the Indian red water
lily be referred to as Nymphaea x rubra, nomen- Nymphaea x rubra is a perennial herbaceous
claturally signifying it to be a natural hybrid. aquatic plant with pinkish, corm-shaped rhi-
Observations from previous cytological (Gupta zomes rooted in the sediment and producing
1980) and reproductive (Mitra and Subramanyam slender stolons. Leaves are simple, orbicular with
1982; Venu et al. 2003) studies also supported the dentate or serrated margin, floating, 20–50 cm in
hybrid origin of N. rubra. Mitra and Subramanyam diameter, sub-peltate, > 0.5 cm from base of
(1982) reported the N. rubra to be propagated sinus, reddish purple and pubescent on abaxial
vegetatively and failed to set fruit in nature. As a surface and glossy lime green or reddish tint on
result of its failure to set fruit/seed, they ques- adaxial surfaces, on long petioles arising from
tioned the treatment of N. rubra as a true species the submerged rhizomes. Submerged leaves are
and rather suggested it as an apomict evidence cordate or sagittate with reddish tint. The flower
strongly pointed to the origin of N. rubra through is showy, fragrant, bisexual, actinomorphic, poly-
hybridization or more appropriately an allopoly- merous, hypogynous up to 15 cm across and soli-
ploidization event involving N. pubescens and tary on emergent, dark-red peduncle above water.
N. lotus followed by chromosome duplication The calyx is aposepalous with four red oblong–
thereby explaining the difference in the ploidy lanceolate, caducous sepals. The corolla is
level of this plant species. apopetalous; 20–25 mm, elliptic to oblanceolate;
its red to purple petals have length thrice longer
than breadth, and many are seriately and spirally
Agroecology arranged (Plate 1). The androecium is polyan-
drous, stamens are 40 to > 50, many are seriately
Nymphaea x rubra is a hydrophyte, found in riv- and spirally arranged, the filaments are petaloid
ers and ponds in still or slow-flowing fresh waters and dark-red purple, and anthers are cream
in tropical and subtropical areas. It thrives in full coloured. The ovary is globose, enveloped in a
sun and water of pH 6–8. fleshy torus, multi-carpellate, syncarpous and
multi-loculed. The fruit, if formed, is a globose
berry containing numerous ellipsoid seeds.
Edible Plant Parts and Uses

The flowers of the plant are used for ornamental Nutritive/Medicinal Properties
purposes, while rhizomes, young leaves and
peduncle are used as food and vegetable (Devi The proximate nutrient content of various parts
et al. 2015). The species is gaining an importance of N. rubra plant had been determined and found
in the local markets of Northeast India as food to be quite high (Devi et al. 2015). The maximum
material. amount of protein was recorded as 23.88 % of
Rhizome is used as vegetable (Pathak and total dry weight. Proximate nutrient composition
Sarma 2013), the tuber is eaten as a vegetable (per 100 g) of edible tuber was reported as: mois-
(Mohan and Kalidass 2010). The young pedun- ture 81.33 %, crude protein 8.31 g, crude lipid
cles are eaten as vegetable. In India, the fried 5.05 g, crude fibre 3.74 g, ash 3.80 g, N-free
seeds are eaten as puffed grains. The petioles, extractives 79.09 %, energy 1650 kJ/100 g, Na
fruit and roots are cooked as vegetables; seeds are 34.1 mg, K 734 mg, Ca 354 mg, Mg 104 mg, P
eaten raw or roasted; fruit and rootstock are eaten 76.3 mg, Zn 1.64mg, Mn 1.34 mg, Fe 28.14 mg,
raw (Pegu et al. 2013; Patiri and Borah 2007). Cu 1.12 mg, starch 13.52 g, niacin 5.88 mg and
120 Nymphaeaceae

Plate 1 Flowers and leaves

of red water lily

ascorbic acid 14.43 mg (Mohan and Kalidass Antidiabetic Activity

2010). Tuber was reported to have an in-vitro
protein digestibility of 5.78 % and in-vitro starch The ethanolic extract of dried N. rubra flowers
digestibility of 18.23 %. Antinutritional factors in and one of its fraction, i.e. chloroform, showed
the edible tubers per 100 g comprised total free significant antihyperglycemic activity on strepto-
phenolics 0.2 g, tannins 0.11 g, hydrogen cyanide zotocin (STZ)-induced diabetic rats, neonatally
0.04 mg, total oxalate 0.42 g, amylase inhibitor STZ-treated rats and high-fructose-, high-fat-fed
1.01 AIU/mg soluble starch and 0.56 trypsin insulin-resistant rats (Rahuja et al. 2013). They
inhibitor TIU/mg protein. Tubers were reported lowered blood glucose level of STZ-induced dia-
to be soaked to remove antinutrients and cooked betic rats and improved glucose tolerance post-
before consumption (Mohan and Kalidass 2010). sucrose load in normal rats. The chloroform
Polysaccharides were purified from N. rubra fraction was also observed to enhance 2-deoxy-
carpels and found to have a degree of polymerisa- (3H) glucose uptake by skeletal muscle cells
tion (DP) value of 359.8 kDa (Cheng et al. 2012). (L-6) in concentration-dependent manner. It also
The hydrolyzed polysaccharides had 51.3 % total inhibited the process of adipogenesis and
sugar content, 2.35 % reducing sugar and total increased the expression of glucose transporter
protein content of 5.28 %. protein-4 (GLUT-4), insulin receptor substrate
(IRS) and phosphatidyl inositol-3-kinase
(PI-3 K). The ethanol extract of Nymphaea rubra
Antioxidant Activity flowers ameliorated TNF-α-induced insulin
resistance in the rat skeletal muscle cell line (L6
Among different concentration tested, methanol myotube) (Gautam et al. 2014). It enhanced the
extract of N. rubra rhizome at 1000 μg/mL exhibited GLUT4-mediated glucose transport in a dose-
potent scavenging activity in all models, viz. DPPH, dependent manner and also increases the tyrosine
hydroxyl radical, superoxide and ABTS radical cat- phosphorylation of both IR-β and IRS-1 and the
ion scavenging activity and reducing power (Daffodil IRS-1-associated PI-3 kinase activity in TNF-α-
and Mohan 2014). The total phenolics and flavo- treated L6 myotubes. It was concluded that N.
noids in the methanol extract were found to be rubra reversed the insulin resistance by the inhi-
0.36 g/100 g and 0.67 g/100 g, respectively. bition of c-jun NH2-terminal kinase and nuclear
Nymphaea x rubra 121

Immunomodulatory Activity Comments

Studies showed N. rubra polysaccharides (NR- The plant is propagated vegetatively by division
PS) to have immunomodulatory effect affecting of rhizome.
the maturation and function of rat dendritic cells
derived from rat bone marrow haematopoietic
cells (Cheng et al. 2012). NR-PS exhibited stimu- Selected References
latory effects on rat dendritic cells and promoted
the secretion of TH1 cytokines. Behera SK, Mohapatra TK, Dash V (2010) Anthelmintic
activity of rhizomes of Nymphaea rubra Linn. Anc Sci
Life 29(3):33–36
Cheng JH, Lee SY, Lien YY, Lee MS, Sheu SC (2012)
Anthelmintic Activity Immunomodulating activity of Nymphaea rubra
Roxb. extracts: activation of rat dendritic cells and
The petroleum ether, methanol and chloroform improvement of the TH1 immune response. Int J Mol
Sci 13(9):10722–10735
extract of N. rubra rhizome exhibited dose- Conrad HS (1905) The waterlilies: a monograph of the
dependent anthelmintic activity against test genus Nymphaea. Carnegie Institution of Washington,
earthworm, Pheretima posthuma (Behera et al. Publication, Washington. 4. 279 pp
2010). The methanolic extract showed remark- Daffodil ED, Mohan VR (2014) In vitro antioxidant activ-
ity of Nymphaea rubra L. rhizome. World J Pharm Res
able anthelmintic activity; at a dose of 5 mg/ml, it 3(4):2178–2189
took 28 min for paralysis and 62 min for worm Devi SA, Thongam B, Handique PJ (2015) Nymphaea
death; at 10 mg/ml it took 17 min for paralysis rubra Roxb. ex Andrews cultivated as an ornamental,
and 51 min for worm death; at 20 mg/ml it took food and vegetable in the North Eastern region of
India. Genet Resour Crop Evol 62:315–320
8 min for paralysis and 27 min for worm death. Dkhar J, Kumaria S, Rao SR, Tandon P (2013) New
The standard albendazole at 10 mg/ml took insights into character evolution, hybridization and
38 min for paralysis and 64 min for worm death. diversity of Indian Nymphaea (Nymphaeaceae): evi-
dence from molecular and morphological data. Syst
Biodivers 11(1):77–86
Gautam S, Rahuja N, Ishrat N, Asthana RK, Mishra DK,
Traditional Medicinal Uses Maurya R, Jain SK, Srivastava AK (2014) Nymphaea
rubra ameliorates TNF-α-induced insulin resistance
The flower is astringent and cardiac tonic and is via suppression of c-jun NH2-terminal kinase and
nuclear factor-κB in the rat skeletal muscle cells. Appl
employed in palpitations of the heart, while the rhi- Biochem Biotechnol 174(7):2446–2457
zomes are used for dysentery and dyspepsia (Khare Gupta PP (1980) Cytogenetics of aquatic ornamentals
2007). Roots are administered for vomiting by rural VI. Evolutionary trends and relationships in the genus
people of Nagaon district, Assam (Sarma and Saikia Nymphaea. Cytologia 45:307–314
Guruge S, Yakandawala D, Yakandawala K (2014)
2010). The Tai-Khamyang of Assam pulverised the Nymphaea rubra Roxb. ex Andrews in Sri Lankan fresh
dried roots to powder and applied the powder in dis- waters. In: Proceedings of international forestry and
eased area for 6–7 days for piles (Sonowal and environment symposium, Sri Lanka. Published by
Barua 2011). In Manipur, north-eastern India, Department of Forestry and Environmental Science,
University of Sri Jayewardenepura. vol 18: P127 (abstract)
crushed rhizome powder is mixed with honey and Juffe Bignoli D (2013) Nymphaea rubra. The IUCN red
used as therapy for bleeding nose, piles and dysen- list of threatened species. Version 2014.2. www.iuc-
tery and as cardiotonic (Yumnam et al. 2012).
Khare CP (2007) Indian medicinal plants: an illustrated
dictionary. Springer, Heidelberg. 900 pp
La-ongsri W, Trisonthi C, Balslev H (2009) A synopsis of
Other Uses Thai Nymphaeaceae. Nordic J Bot 27(2):97–114
Mitra RL, Subramanyam K (1982) Is Nymphaea rubra
Red water lily is used as an ornamental plant in Roxb. ex. Andrews an apomict? Bull Bot Surv India
aquatic landscaping and also in the aquarium Mohan VR, Kalidass C (2010) Nutritional and antinutri-
trade. The cut flowers are used for religious pur- tional evaluation of some unconventional wild edible
poses in India. plants. Trop Subtrop Agrosyst 12:495–506
122 Nymphaeaceae

Pathak U, Sarma DC (2013) A critical study of the biore- Sarma SK, Saikia M (2010) Utilization of wetland
sources of Khaamronga Beel – a wetland for sustain- resources by the rural people of Nagaon district
able development for the people of Thakurkuchi Assam. Indian J Trad Kwon 9(1):145–151
village of Chandrapur area of Assam. Indian J Fund Sonowal R, Barua I (2011) Ethnomedical practices among
Appl Life Sci 3(2):245–251 the Tai-Khamyangs of Assam, India. Ethno Med
Patiri B, Borah A (2007) Wild edible plants of Assam. 5(1):41–50
Geetakhi Printers and Publishers, Guwahati USDA-ARS (2015) National genetic resources program.
Pegu R, Gogoi J, Tamuli AK, Teron R (2013) Germplasm Resources Information Network – (GRIN)
Ethnobotanical study of wild edible plants in Poba [Online Database]. National Germplasm Resources
reserved forest, Assam, India: multiple functions and Laboratory, Beltsville, Maryland. https://npgsweb.ars-
implications for conservation. Res J Agric For Sci .
1(3):1–10 Accessed 7 Mar 2015
Rahuja N, Mishra A, Gautam S, Tamrakar AK, Maurya R, Venu P, Rajakumar TS, Danile P (2003) On the identity of
Jain SK, Srivastava AK (2013) Antidiabetic activity in and sterility in Nymphaea rubra Roxb. ex Andrews
flowers of Nymphaea rubra. Int J Pharm Sci Rev Res (Nymphaeaceae). Rheedia 13:29–34
22(24):121–133 Yumnam RS, Dev CO, Abujam SKS, Chetia D (2012)
Study on the ethnomedicinal system of Manipur. Int
J Pharm Biol Arch 3(3):587–591
Victoria amazonica

Scientific Name Victória-Regia (Portuguese) Irupé (Guarani),

Uapé, Aguapé (Tupi)
Victoria amazonica (Poepp.) Sowerby Chinese: Wang Lin
Czech: Viktorie Amazonská, viktorie královská
Estonian: Amasoonase Viktooria
Synonyms French: Victoria d’Amazonie
German: Amazonas-Riesenseerose, Amazonas
Euryale amazonica Poepp. (basionym); Wasserlilie, Königliche Seerose, Viktoria
Nymphaea victoria R. H. Schomb. ex Lindl., Lithuanian: Gigantiškoji viktorija
nom. inval.; Victoria amazonica Planch. ex Casp.; Polish: Wiktoria Królewska
Victoria regia Lindl.; Victoria regia var. randii Spanish: Abatiyú, Agoapé, Aguapé, Hoja De
hort. ex Conard, nom. inval.; Victoria regina Sol, Iguapé, Irupé, Maíz De Agua, Maruru,
R. H. Schomb Ninfa Real, Plato De Agua, Loto Gigante
Swedish: Jättenäckros
Upper Sorbian: Wodowa krasnica


Common/English Names The species is native to the shallow waters of the

lakes and waterways in the Amazon River Basin,
Amazon Water Lily, Amazon Water Platter, Giant the Guianas and the Pantanal in South America.
Amazon Water Lily, Giant Water Lily, Royal Its aquatic tropical home spans Brazil, Colombia,
Water Lily, Victoria Lily, Water Maize Peru and Guyana.

Vernacular Names Agroecology

Brazil: Vitória-Regia, Aguapé-Assú, Cará- Victoria amazonica is tropical aquatic. The plant
D’água, Forno-D’água, Forno-De-Jaçanã, roots are best anchored in organically rich loams
Jaçanã, Milho-D’água, Mururê, Nanpé, at the bottom of the bayous. Plants will remain
Rainha-Dos-Lagos, Rainha-Dos-Nenúfare, perennial as long as water temperatures remain at
© Springer International Publishing Switzerland 2016 123
T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_9
124 Nymphaeaceae

least 24 °C and thrive in calm shallow waters tate, floating, entire and full green adaxially with
(<2 m), full sun, slightly alkaline to neutral the edge turn upwards forming a rim (5–10 cm
pH. Pollination by scarab beetles happens at high), and the basal surface is a coppery colour
night. with a prominent network of flattened ribs radiat-
ing from the petiole and cross ribs, all bearing
sharp prickles (Plates 1–3) (Anonymous 1850;
Edible Plant Parts and Uses Allen 1854). Kunii et al. (2006) reported that the
maximum value in leaf diameter was 2.35 m
In the Brazilian Amazon, Victoria amazonica (4.34 m2 in area) with a mean value of 1.6–1.7 m.
seeds and rhizomes are important items con- Each plant has 10–15 floating leaves and the total
sumed by riverine inhabitants (Piedade and Junk leaf area per plant differed considerably (mini-
2000). The root, stem and seed of the Victoria lily mum value 12.1–17.2 m2, maximum value 27.6–
are all edible, and the species is also used in tra- 31.2 m2). Flower solitary on an aculeate, terete
ditional medicine (Castner et al. 1998). pedicel arising from the rhizome, longer than leaf
The root, stem and seeds are edible. Dried petiole, pear-shaped in bud (Plates 2–3). The
seeds taste like popcorn. Indigenous Brazilians flower is epigynous, fragrant, 30–50 cm across,
make flour from the seeds (Les 2003). hemicyclic, dichlamydeous, bisexual with ves-
pertine anthesis (Rosa-Osman et al. 2011) and
pollinated by scarab beetles. The calyx, consist-
Botany ing of four large and four small spiny sepals
(Kunii et al. 2006), is coriaceous, fleshy and
Victoria amazonica is a rhizomatous aquatic, green becoming reddish purple. Petals are numer-
strongly aculeate, herbaceous perennial (Plate 1) ous, inserted on the throat and torus of the calyx;
with short, thick, tuberous rhizome, which is outer ones are completely reflexed. The petals
brown externally and white inside and turning remained expanded, erect and white all night and
purple when cut; rhizome bears numerous cylin- the next morning; by the evening of the second
drical adventitious roots abounding in air cavi- day, the petals had turned variegated white pink
ties. Leaves are borne on a submerged aculeate, or red. Stamens are numerous, inserted with the
terete petiole 6 m long radiating from the sub- petals on the torus in about three rows; the sta-
merged rhizome; they are huge, orbicular, pel- mens, including the outer staminodia (infertile

Plate 1 Aquatic, rhizomatous

plant with seven or more large,
orbicular leaves with upturned
Victoria amazonica 125

Plate 2 Huge leaves at the apex of

long radiating submerged petioles
arising from the rhizome

campanulate at the top, with many locules con-

taining numerous anatropous ovule; style is
wanting, and stigmas are forming radiating lines
on the top of the ovary. The fruit is fleshy, cam-
panulate and irregular dehiscent with pseudo-
syncarpy (Rosa-Osman et al. 2011), prickly
11 cm across. Seeds are many and ellipsoid glo-
bose with horny testa and aril that acts in the
water dispersion.

Nutritive/Medicinal Properties
Plate 3 Young leaf with coppery basal surface and
flower bud enclosed by four reddish-purple, spiny sepals Two anthocyanins were isolated from Victoria
amazonica leaves and identified as delphinidin
3-O-(2″-O-galloyl-β-d-galactopyranoside) and
stamens) petaloid, middle fertile stamens and cyanidin 3-O-(2″-O-galloyl-β-d-galactopyranoside)
inner staminodia (termed paracarpels by earlier (Strack et al. 1992). Four esters, together with
authors), were erect and formed an approx. free benzyl alcohol, were found as the main
12-mm tunnel into the floral chamber (Seymour components of the fragrance of Victoria (Kite
and Matthews 2006). The filaments are subulate, et al. 1991). The compounds were methyl
erect, outer filaments petaloid, connective long- 2-methylbutyrate, methyl tiglate, benzyl 2-meth-
appendiculate, filaments of the sterile stamens of ylbutyrate and benzyl tiglate. A number of other
the innermost whorl with fleshy appendages; methyl and benzyl esters were found but at much
anthers introrse. Carpels are numerous, crescent lower concentrations. Fifteen compounds were
or horseshoe shaped up to the time of anthesis, isolated from the methanol leaf extract including
and then postgenital fusion occurs forming the stigmasterol, vanillic acids and aristophyll
ovary. The ovary is globose below and concave (Chang et al. 2014).
126 Nymphaeaceae

Na, K, Rb, Cs, AG, Zn, Y, Nb, Se, F, Br, Cl, I Comments
and Ni increased with age; Li, Cu, Be, Mg, Sr,
Cd, Hg, Ce, Si, Ti, Zr, V, Cr, Fe and Al decreased The plant is depicted in the Guyanese coat of
with age; and Ba, B, Ga, Sc, La, P, Bi and S arms.
exhibited no consistent relationship to various Protogyny has been studied in Victoria ama-
growth stages (Cowgill and Prance 1982). zonica (Prance 1974, 1980). The flowers will
However, Ca, Sn, Pb, As, Mo, Mn and Co were open following a dark period of 30 min, begun
low in concentration in young plants, increased not earlier than 4 p.m. (Gessner 1960). The plants
in the pre-flowering stage and then declined with are often pollinated by scarab beetles (Prance and
the stage of the plant. V. amazonica accumulated Arius 1975; Prance 1980).
Na in excess of that encountered in its terrestrial
relatives. It contained rare earths lanthanum (La)
and cerium (Ce) in normal amounts. Rare earths Selected References
of higher atomic numbers were not detectable.
Mg, Ca, Al, Si, S, Fe, Mn, Li and Ce were signifi- Allen JF (1854) Victoria regia or the great water liliy of
cantly more concentrated in the damaged leaf
sections than in undamaged leaves of the mature Anonymous (1850) Victoria regia. The Gardeners’
plants with unopened buds (Cowgill and Prance Chronicle, pp. unknown
1989). In the case of pre-flowering plants, Na, Castner JL, Timme SL, Duke JA (1998) A field guide to
medicinal and useful plants of the Upper Amazon.
Mg, Ca, Al, Si, S, Cl, Fe and Mn were more prev-
Feline Press, Gainesville. 154 pp
alent in the damaged tissue than in the unattacked Chang MY, Wu HM, Li HT, Li WJ, Chen SJ, Chen CY
leaves of the same plants. In the former case, Na, (2014) Secondary metabolites from the leaves of
K, Cl and P are rapidly lost when damage occurs, Victoria amazonica. Chem Nat Compd
while plants that have had a chance to recover
Cowgill UM, Prance GT (1982) Changes in the chemical
from the damage, as in the case of the pre- composition during growth stages of Victoria amazo-
flowering plant leaves, begin to make up the dif- nica (Poeppig) J. de C. Sowerby (V. regia auctt.),
ference in elemental concentration between the Nymphaeaceae. Int Rev Gesamten Hydrobiol
damaged and undamaged leaves. Mineralogical
Cowgill UM, Prance GT (1989) A Comparison of the
investigation showed that the amount of calcite chemical composition of injured leaves in contrast to
(CaCO3), the calcium oxalates weddellite and uninjured leaves of Victoria amazonica
whewellite and the siliceous minerals – low-form (Nymphaeaceae). Ann Bot 64(6):697–706
Duke JA, Vásquez R, Vásquez VM (1994) Amazonian
cristobalite opal, low-form tridymite opal and
ethnobotanical dictionary. CRC Press, Boca Raton,
quartz – was higher in the recently injured leaves 215 pp
than in those that had been injured some time pre- Gessner F (1960) Die Blütenöffnung der Victoria regia in
viously or than in the uninjured leaves. ihrer Beziehung zum Licht. Die Blütenöffnung der
Victoria regia in ihrer Beziehung zum Licht. The
The plant extract has folkloric medicinal use
opening of flowers of Victoria regia in relation to light.
for rheumatism, inflammation and haemorrhoids Planta 54:453–463 (In German)
when mixed with andiroba oil (Duke et al. 1994). Huxley AJ, Griffiths M, Levy M (eds) (1992) The new
RHS dictionary of gardening, (4 vols). MacMillan,
Kite G, Reynolds T, Prance GT (1991) Potential
Other Uses pollinator-attracting chemicals from Victoria
(Nymphaeaceae). Biochem Syst Ecol 19(7):535–539
V. amazonica is used as an aquatic ornamental in Kunii H, Sunanisari S, Fukuhara H, Nakajima T, Widjaja
F (2006) Leaf expansion rate and life span of floating
botanical gardens and parks.
leaves in Victoria amazonica (Poepp.) Sowerby grow-
ing in Kebun Raya, Bogor, Indonesia. Tropics
Victoria amazonica 127

Les DH (2003) Nymphaeales. In: eLS. John Wiley & Rosa-Osman SMD, Rodrigues R, Mendonça MSD,
Sons Ltd, Chichester. doi:10.1038/ Souza LAD, Piedade MTF (2011) Morfologia da
npg.els.0003697 flor, fruto e plântula de Victoria amazonica (Poepp.)
Piedade MTF, Junk WJ (2000) Sustainable management of J.C. Sowerby (Nymphaeaceae). Acta Amazonica
species diversity and primary production of herbaceous 41(1):21–28. [Morphology of flower, fruit and seed-
plants of the Central Amazon floodplain. German- ling of Victoria amazonica (Poepp.) J.C. Sowerby
Brazilian Workshop on neotropical ecosystems – (Nymphaeaceae)]
Achievements and Prospects of Cooperative Research Schneider EL (1976) The floral anatomy of Victoria
Hamburg. Session 6: concepts and paradigms for man- Schomb. J Linn Soc Bot 72:115–148
agement of ecosystem resources. Hamburg Seymour RS, Matthews PGD (2006) The role of ther-
Prance GT (1974) Victoria amazonica ou Victoria regia? mogenesis in the pollination biology of the amazon
Acta Amazon 4(3):6 waterlily Victoria amazonica. Ann Bot 98:
Prance GT (1980) A note on the pollination of Nymphaea 1129–1135
amazonum Mart. & Zucc. (Nymphaeaceae). Brittonia Strack D, Wary V, Metzger JW, Grosse W (1992) Two
32(4):505–507 anthocyanins acylated with gallic acid from the leaves
Prance GT, Arius JR (1975) A study of the floral biology of Victoria amazonica. Phytochemistry 31(3):
of Victoria amazonica (Poepp.) Sowerby 989–991
(Nymphaeaceae). Acta Amazon 5(2):109–139
Diuris semilunulata

Scientific Name Agroecology

Diuris semilunulata Messmer This orchid grows on well-drained, moist soils

among grass and shrubs in sclerophyll forest;
south from Nerriga, southern Tablelands and in
Synonyms Victoria.

None recorded
Edible Plant Parts and Uses

Family Starch tubers of Diuris spp. including this species

are edible but bland and glutinous and eaten by
Orchidaceae the aborigines (Cribb and Cribb 1987; Low 1989,

Common/English Names
Donkey Orchid, Late Leopard Orchid, Spotted
Double-Tail Orchid Diuris semilunulata is a terrestrial, perennial,
geophytic herb. Roots are filamentose with small,
naked, ovoid or cylindrical, often paired, tubers.
Vernacular Names It has 15–25 cm long and 3–4 mm wide, linear,
basal, grass-like, alternate or whorled leaves, that
None recorded is, conduplicate, with 2 leaves in a tuft (Plate 1).
Scape is 20–35 cm high with 3–5-flowered, loose
raceme. Flowers are hermaphroditic, orange,
Origin/Distribution heavily blotched and suffused with brown and
purple. Dorsal sepal is ovate, 6–10 mm long,
This orchid is native to Australia. 7–10 mm wide and erect. Lateral sepals are linear

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Diuris semilunulata 129

to oblanceolate, 12–18 mm long, 2–4 mm wide,

crossed and recurved beneath the labellum. Petals
are divergent, ovate to obovate and 5–11 mm
long by 4–9 mm wide; claw is 5–9 mm long.
Labellum is 4–8 mm long; lateral lobes are nar-
rowly to broadly, cuneate, 5–8 mm long and
2–7.5 mm wide; margins are crenulate; mid-lobe
is cuneate, 5–7 mm wide and ridged along mid-
line; and callus is with two 3–6 mm long raised
ridges. Its fruit has a non-fleshy, dehiscent cap-
sule, containing between 30 and 500 minute
light-dark coloured, winged seeds.

Nutritive/Medicinal Properties

No information on its nutritive value and medici-

nal properties had been published.

Other Uses

This orchid is mainly grown by orchid enthusi-

asts and some small commercial orchid growers.
Plate 1 Flowers of Diuris semilunulata

Jones DL (1970) Some thoughts on the taxonomy of the

Comments genus Diuris. Orchadian 3:119–120
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
New South Wales, vol 3. New South Wales University
This orchid is propagated from seeds and from Press, Kensington. 775 pp
the underground tubers. Low T (1989) Bush Tucker – Australia’s wild food har-
vest. Angus & Robertson, North Ryde, 233 pp
Low T (1991) Wild food plants of Australia. Angus &
Robertson, North Ryde. 240 pp
Selected References Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN (eds)
(2001) Genera Orchidacearum: Volume 2 Orchidoideae
Cribb AB, Cribb JW (1987) Wild food in Australia, 2nd (Part one). Oxford University Press, Oxford/New York,
edn. Fontana Collins, Sydney. 240 pp 438 pp
Diuris sulphurea

Scientific Name Origin/Distribution

Diuris sulphurea R.Br. This orchid is native to Australia.

Synonyms Agroecology

Diuris latifolia Rupp., Diuris oculata F. Muell. This orchid is commonly found in well-drained
ex Lindl., Diuris sulphurea f. immaculata sul- moist soil in semi-shaded sites on slopes of the
phurea Gand., Diuris sulphurea f. tasmanica foothills in open sclerophyll forest and health in
Gand. South Australia, Tasmania, Victoria, New South
Wales and Queensland.

Edible Plant Parts and Uses
Starchy root tubers are edible but bland and gluti-
nous and eaten by the aborigines (Anonymous 2010,
Common/English Names 2011; Cribb and Cribb 1987; Low 1989, 1991).

Hornet Orchid, Tiger Orchid, Yellow Tiger

Orchid Tiger Orchid, Hornet Orchid Botany

Diuris sulphurea is a terrestrial, perennial, geo-

Vernacular Names phytic herb. Roots are filamentose with small,
naked, ovoid or cylindrical, often paired, tubers.
None recorded It has 15–25 cm long and 3–4 mm wide, linear,
basal, grass-like, alternate or whorled leaves, that
is, conduplicate, with 1–3 leaves in a tuft (Plates 1

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Diuris sulphurea 131

and 2). Scape is 30–60 cm high with 1–6-flowered

racemose inflorescence. Its flowers are bright
yellow with prominent dark markings on the
labellum and dorsal sepal, 3–4 cm across and
stalked (Plate 3). Dorsal sepal is ovate, 8–20 mm
long, 7–15 mm wide and obliquely erect. Lateral
sepals are linear to lanceolate, 10–25 mm long,
1–3 mm wide, deflexed, parallel or crossed.
Petals are divergent, ovate, 5–18 mm long and
5–12 mm wide; apex is obtuse or notched; and
claw is 3–10 mm long and dark coloured.
Labellum is 7–15 mm long; lateral lobes are
broad cuneate, 3–6 mm long and 2.5–4.5 mm
wide; margins are entire or crenulate; and callus
is with 3–5 mm long single ridge. Column is with
stamen and style nearly free; anther is dorsal and
basifixed; stigma has large scutiform; and ovary
is elongate, ribbed and glabrous. Capsule is
dehiscent, thin walled and glabrous erect contain-
ing numerous light-dark-coloured, winged seeds.

Nutritive/Medicinal Properties

No information on its nutritive value and medici- Plate 1 Long scape with a few flowered raceme
nal properties had been published.

Plate 2 Basal linear, grass-like

conduplicate leaves
132 Orchidaceae

Other Uses

This orchid is mainly grown by orchid enthusi-

asts and some small commercial orchid growers.


This orchid is propagated from seeds and from

the underground tubers.

Selected References
Anonymous (2010) Diuris sulphurea. http://www.yarra-
Anonymous (2011) Diuris sulphurea (Tiger orchid).
h t t p : / / bu s h c r a f t o z . c o m / f o r u m s / s h ow t h r e a d .
Cribb AB, Cribb JW (1987) Wild food in Australia, 2nd
edn. Fontana Collins, Sydney, 240 pp
Jones DL (1970) Some thoughts on the taxonomy of the
genus Diuris. Orchadian 3:119–120
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
New South Wales, vol 3. New South Wales University
Press, Kensington. 775 pp
Plate 3 Bight yellow flowers with prominent dark- Low T (1989) Bush Tucker – Australia’s wild food har-
coloured markings vest. Angus & Robertson, North Ryde, 233pp.
Low T (1991) Wild food plants of Australia. Angus &
Robertson, North Ryde. 240 pp
Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN (eds)
(2001) Genera Orchidacearum: Volume 2 Orchidoi-
deae (Part one). Oxford University Press, Oxford/
New York, 438 pp
Pterostylis curta

Scientific Name Agroecology

Pterostylis curta R.Br. The species is commonly found in sclerophyll

forest in coastal southeast Queensland, New
South Wales, Victoria, southeast South Australia,
Synonyms Tasmania and Lord Howe Island. The species
prefer moist and well-drained sites in the wild,
None recorded but the range of forest types they occur in is
broad, from grassy woodlands to riparian forests
to subtropical rainforest, and even in exotic pine
Family plantations. They occur in semi-shade to full

Edible Plant Parts and Uses

Common/English Names
The tubers are edible and eaten by the aborigines
Blunt Greenhood (Anonymous 2010; Steenbeeke 2001).

Vernacular Names Botany

None recorded Pterostylis curta is a terrestrial perennial herb,

10–30 cm high, arising from small round tubers
to form large colonies. Leaves are in ground-
Origin/Distribution hugging rosette of 2–6, shortly petiolate, dark
green, ovate to elliptic or oblong and 1.5–10 cm
The species is indigenous to Australia. long by 8–30 mm wide, with entire or wavy mar-

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DOI 10.1007/978-3-319-26062-4_12
134 Orchidaceae

Plate 1 Pterostylis curta with ground-hugging rosette of

leaves Plate 2 Long scape with single flower

Plate 3 Close up of flowers

Pterostylis curta 135

gins (Plates 1 and 2). Scape is 30 cm with a single Other Uses

erect flower, that is, 35 mm long, white and green
with brown suffusions in the galea. Central sepal This orchid is mainly grown by orchid enthusi-
and petals are united into a galea (hood) over the asts and some small commercial orchid growers.
column and labellum (lip) (Plates 2 and 3). Galea
is mostly upright; top, shortly curved; and tip,
tinged green brown and pointed. Lateral sepals Comments
are erect, loosely embracing the hood, leaving a
broad V-shaped lateral gap; free points are linear The plant is readily propagated from the tubers.
tapered, about10 mm long, divergent and
recurved. Labellum is oblong-obovate, brown
and twisted. Selected References
Anonymous (2010) Pterostylis curta. http://www.yarra-
Nutritive/Medicinal Properties Directory/Lower_Storey/Orchids/Pterostylis_curta
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
No information on its nutritive value and medici- New South Wales, vol 3. New South Wales University
nal properties had been published. Press, Kensington. 775 pp
Steenbeeke G (2001) List of plant species from northern
NSW that may be used as food plants. http://brg.cma.
Pterostylis pedunculata

Scientific Name Agroecology

Pterostylis pedunculata R. Br. This orchid occurs in moist sheltered habitats in

sclerophyll forest of coastal and near-coastal dis-
tricts, in Queensland, New South Wales, Victoria,
Synonyms Tasmania and Lord Howe Island. It thrives on
moist soil in leaf litter in fern gullies and moist to
None recorded wet forests in dappled to full shade.

Family Edible Plant Parts and Uses

Orchidaceae The starchy tubers are edible and eaten by aborig-

ines (Steenbeeke 2001; Anonymous 2011).

Common/English Names
Pterostylis pedunculata is a terrestrial perennial
herb, 25 cm high, arising from small round
Vernacular Names tubers to form large colonies. It has a ground-
hugging rosette of 2–6, shortly petiolate, thick,
None recorded dark green, ovate to oblong, 10–65 mm long by
6–18 mm wide leaves, with entire or wrinkled
margins (Plates 1 and 2). Scape is 20–25 cm
Origin/Distribution with a single erect flower, that is, 15 mm long,
white and green with reddish-brown to black
The species is indigenous to Australia. suffusions in the galea. Central sepal and petals

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Pterostylis pedunculata 137

Plate 1 Pterostylis pedunculata

plant habit

Plate 2 Ground-hugging rosette of


are united into a galea (hood) over the column wide, dark reddish brown, straight and apex
and labellum (lip) (Plates 2, 3 and 4). Apex of obtuse.
galea is horizontal. Dorsal sepal is acute to sub-
acute. Lateral sepals are erect, loosely embrac-
ing the hood, leaving a broad V-shaped lateral Nutritive/Medicinal Properties
gap; free points are filiform, 15–30 mm long,
divergent and recurved. Petals are narrow and No information on its nutritive value and medici-
acute. Labellum is ovate, 4–7 mm long by 3 mm nal properties had been published.
138 Orchidaceae

Plate 3 Scape with single


Other Uses

This orchid is mainly grown by orchid enthusi-

asts and some small commercial orchid growers
as ornamental pot orchids.


The plant is readily propagated from the small


Selected References
Anonymous (2011) Pterostylis pedunculata. http://www.
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
New South Wales, vol 3. New South Wales University
Press, Kensington. 775 pp
Steenbeeke G (2001) List of plant species from northern
NSW that may be used as food plants. http://brg.cma.

Plate 4 Close-up of flower

Oxalis tuberosa

Scientific Name Bolivia: Apiña, Apilla

Brazil: Azedinha, Batata-Baroa, Mandioquinha
Oxalis tuberosa Molina Chile: Cubia
Columbia: Hibia, Huasisai, Ibi, Ibia, Ibiasoca
Danish: Oka
Synonyms Dutch: Peruaanse Klaverzuring
Ecuador: Oca
Acetosella crenata (Jacq.) Kuntze, Acetosella Finnish: Oka
tuberosa (Molina) Kuntze, Oxalis arracatcha hort. French: Oca D’amérique, Oxalide Crénelée,
ex Zucc., Oxalis arracacha G. Don, Oxalis chicli- Oxalide Tubéreuse, Oxalis Tubéreux, Surelle
gastensis R. Knuth, Oxalis crassicaulis Zucc., Tubéreuse, Truffette Acide
Oxalis crenata Jacq., Oxalis melilotoides var. German: Knollen-Sauerklee, Oka, Peruanischer
argentina Griseb., Xanthoxalis crassicaulis (Zucc.) Sauerklee
Small, Xanthoxalis tuberosa (Molina) Holub Italian: Ossalide Crenata, Trifoglio Tuberose
Mexico: Papa Colorada, Papa Extranjera, Papa Roja
New Zealand: Yam
Family Peru: Apiña, Oca, Oqa
Polynesia: Yam
Oxalidaceae Quechuan: O’qa, Okka, Uqa
Spanish: Aleluya Tuberosa, Cubio, Ibia, Oca
Swedish: Oca
Common/English Names Turkish: Yabani Kuzu Kulagi
Venezuela: Cuiba, Ciuva, Macachin, Miquichi,
Kao, Oca, Oka, New Zealand Yam, Papa Roja, Oxalida, Quiba, Ruba, Timbo
Quiba Yam

Vernacular Names Origin/Distribution

Arabic: Hhummâd Oxalis tuberosa is indigenous to the Andes in

Argentina: Miquichi South America (Arbizu and Tapia 1994). The
Aymara: Apiña, Apilla, Kawi area of greatest diversity both of cultivated and

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DOI 10.1007/978-3-319-26062-4_14
140 Oxalidaceae

wild forms is found in the central region of Peru thrives best in light, well-drained and rich soil in
(latitude 10°S) and northern Bolivia (latitude full sun. Oca requires day length shorter than
20°S). Oca is grown in greatest abundance in the 12 h to stimulate tuber formation; longer day
highlands of Ecuador, Peru, Columbia and length periods promote foliage growth.
Bolivia, although it is found as far north as
Venezuela and as far south as Chiloe Island in
Chile (Popenoe et al. 1989). Of all Andean root Edible Plant Parts and Uses
and tuber crops, oca is presently second only to
potato in area planted within the Central Andean Oca tubers are boiled, baked, roasted, fried,
region. Oca was introduced into Europe in the mixed fresh with salads, added to stews and
last century but did not establish as a permanent soups and pickled in vinegar or served as a sweet,
vegetable crop. In recent decades, oca has estab- either plain or candied (Arbizo and Tapia 1994;
lished as a popular commercial vegetable crop King and Gerhoff 1987; Popenoe et al. 1989;
in New Zealand, where it called yam (Popenoe Bradbury and Emshwiller 2011). In the Andes, a
et al. 1989). few genotypes types are eaten raw. In Mexico,
Earlier results of studies with nuclear-encoded, oca is commonly sprinkled with salt, lemon and
chloroplast-expressed glutamine synthetase sug- hot pepper, and eaten raw. It is also made into
gested that Oxalis picchensis might be a progeni- bottled preserves (often in vinegar). Ocas that are
tor of oca (Emshwiller and Doyle 2002). consumed fresh belong to the category wayk’u
Subsequent AFLP (amplified fragment length (boiling) or misk’i (sweet, delicious). Aymara
polymor phism) data of this species, as well as speakers refer to ocas for processing as luk’i and
different populations of wild, tuber-bearing those for fresh consumption as q’eni. In the
Oxalis found in Lima Department, Peru, were Andes, Oca is first sun-dried for a few days, to
relatively divergent from O. tuberosa (Emshwiller make it sweeter and then parboiled, roasted or
et al. 2009). Results from all analytical methods prepared as pachamanca (meat roasted in a hole
suggested that the unnamed wild, tuber-bearing in the ground). The dried, frozen tuber is called
Oxalis found in Bolivia and O. chicligastensis in khaya. If it is washed after freezing, a whiter
NW Argentina were the best candidates as the product called okhaya is obtained which is con-
genome donors for polyploid O. tuberosa. sidered to be of superior quality. The flour of the
latter is used to make porridges and desserts.
Bitter varieties are almost always converted into
Agroecology dry products (cavi or caya), during which the bit-
terness disappears to leave bland-tasting products
Oca is widely grown as a staple crop in the that can be stored without refrigeration. The bit-
Andean highlands due to its tolerance of mar- terness can also be removed by soaking and
ginal soil, acidities between pH 3.5–7.8, high repeatedly freeze-drying the tubers. Oca tubers
altitude, harsh climate and ease of propagation also have potential for producing superior quality
(Popenoe et al. 1989; Arbizu and Tapia 1994). In starch for the food industry or alcohol.
its native range, it is cultivated from 2800 to Sensory evaluation studies in New Zealand
4000 m altitudes. In New Zealand, it is grown found that for raw oca, panellists preferred bright
near sea-level. Oca flourishes in a climate of low red tubers, but size was also important (Sangketkit
cool temperatures from 4 to 18 °C and mean et al. 2000). Flesh colour, bitterness and meali-
annual precipitation of 570–2000 mm uniformly ness were important variables which described
distributed throughout the growing season. It tol- the overall acceptability of steamed oca, while
erates light frost but its top growth is severely only flesh colour and bitterness were important
damaged by sub-zero temperatures. Temperatures variables for baked oca. Cultivars which had flesh
above about 28 °C cause the plant to wilt and its colour described as ‘bright yellow’, flavour as
leaves to die and tuber production reduced. It ‘no bitter taste’ and a ‘very slightly mealy’ tex-
Oxalis tuberosa 141

ture were more preferred. Cultivars with higher and form tubers. The tubers are claviform-ellip-
yellow/blue (b∗) and chroma (c∗) values for soid and cylindrical, 3–15 cm long by 3 cm wide,
cooked tuber skin and flesh colours were more stubby-looking, with buds on the whole surface,
acceptable. Martin et al. (2005) found that culti- and variegated in colour: white, yellow, red and
vars with red skin, bright yellow flesh colour, a purple. Leaves trifoliate on 2–9 cm long petioles,
slightly mealy texture and a sweet taste were leaflets, heart-shaped, clover-like and green.
preferred. Inflorescence with 4–5 small flowers. Calyx with
five pointed green sepals. Corolla with five
obovate, clawed petals fused at the base and with
Botany radiating purple striations at the basal part.
Stamens ten. Flowers are tristylous: short-styled
Oca is a compact, perennial, tuberous herb, erect flowers with two tiers of stamens above the style;
becoming decumbent or prostrate towards matu- mid-styled flowers with a tier of stamens both
rity, usually 20–30 cm high, with cylindrical, above and below the styles and long-styled flow-
succulent stems that vary in colour from yellow ers with two tiers of stamens below the styles.
and green to a purplish red (Plates 1–2). Under
long days, the stolons grow as above-ground
stems; under short days, they penetrate the soil Nutritive/Medicinal Properties

The nutritional composition (dry weight basis) of

oca tubers was reported as protein 3.0–8.4 %,
carbohydrate 83.0–88.8 %, fat 0.5–0.6 %, ash
1.9–3.5 %, fibre 4.0–5.1 %, moisture 80.2–
84.6 %, energy 368–374 cal/100 g (King and
Gershoff 1987). They also contained all of the
essential amino acids (mg/g protein): lysine
57–59 mg, threonine 44–47 mg, valine 26–48 mg,
isoleucine 36–468 mg, leucine 53–60 mg, phe-
nylalanine + tyrosine 57–69 mg, tryptophan 5.5–
8.0 mg and methionine + cystine 25–34 mg.
Nutrient values (ppm, dry weight basis) of oca
Plate 1 Variously coloured oca tubers (CIP) tubers reported by Duke (1994 onwards): carbo-
hydrate 138–852 ppm, protein 7–62, fat 6–37,
fibre 5–49, ascorbic acid 370–2284, Ca 40–247,
Fe 8–49, P 340–2099, riboflavin 0.3–4.3, thia-
mine 0.5–3.1, b-carotene 0.2, energy 630–3890
The main proteins in tubers of 23 different oca
accessions were quite acidic with isoelectric
points pH 4.7–4.9 and relative molecular weight
of 17–18 KD (Stegemann et al. 1988). The pro-
teins were rich in aspartic and glutamic acid (one
third of all amino acids, asp:glu ratio of 1:2) with
a considerable share of essential ones: histidine
and lysine (7.5–9 %); leucine, isoleucine and
valine (3.5–4 %); threonine (5 %); and arginine
(4.5–7 % in most except 15 % in accession
Plate 2 Harvested pinkish-red oca tubers CTO-140).
142 Oxalidaceae

Oca tuber was reported to have a nutritional Hernández-Lauzardo et al. (2004) reported
value equivalent to or better than potato; on aver- that oca starch possessed an apparent amylose
age, oca contained 84.1 % moisture, 1.1 % protein content of 33 %, similar to maize starch used as
and 0.6 % lipid (Popenoe et al. 1989). Fourteen control, with a granule size between 25 and
percent of its nutrient content comprised carbohy- 50 μm with oval and elliptical shapes and A-type
drates with sucrose as the major sugar (Gross X-ray diffraction pattern. The gelatinization tem-
et al. 1989). Chemical composition of oca tubers perature of oxalis starch was 64.0 °C, that was
contains protein 6.84 %, lipid 0.72 %, soluble lower than the one determined in maize starch
fibre 0.29 %, insoluble fibre 6.85 %, total sugar (73.0 °C), with an enthalpy value of 12.2 J/g,
23.28, starch 56.82 %, mineral residues 4.48 % which was similar to that of maize starch. Both
and moisture 86.23 % (Valcárcel-Yamani et al. oxalis and maize starch pastes behaved as weak
2013). Chemical composition of oca contains viscoelastic systems with the elastic modulus
starch 99.21 %, amylose 27.60 %, mineral resi- (G′) predominating over the viscous character
dues 0.15 % and moisture 10.93 %. Oca starch (G″). Heating (gelatinization) caused a more pro-
granules were ellipsoid, oval, conical, pear-shaped nounced enhancement in the structure of the
and prismatic forms: ellipsoids and oval granules maize starch pastes than in that of oxalis starch
with lengths up to 54.30 μm. The physical, chemi- pastes. They concluded that due to its physico-
cal and functional characterization of starches chemical, functional and rheological properties,
from Andean tubers oca, mashua and ulluco sug- oxalis starch could be suitable for testing its use
gested that these starches could be used in food in the cosmetic and food industry.
systems and other industrial applications, in prod- Total phenolic content of oca tubers ranged
ucts that require easy cooking, hot high viscosity, from 0.71 to 1.32 mg/g with the purple coloured
stability under refrigeration and do not need to be varieties having higher content and the yellow
frozen. The absence of protein in the isolated coloured varieties lower phenolic content
starches indicated the utility of these starches for (Campos et al. 2006). Total carotenoid content
preparing syrups with high glucose content. The ranged from 2 to 25 μg/g in oca, moisture content
starches were found to cook easily and to have a 80–85 % and lipophilic antioxidant in oca tubers
high degree of swelling and solubility, high vis- ranged from 69 to 320 μg/g TE. For oca pig-
cosity, low stability to stirring and cooking or mented tubers, the total anthocyanin content
mechanical action and a low tendency towards ranged from 0.14 to 1.3 mg/g; anthocyanin was
retrogradation. These starches showed high clar- concentrated in the tuber skin.
ity, but with high syneresis when subjected to Eight different anthocyanins were found in the
freeze-thaw cycle. Oca starch granules were oval- coloured tubers of isla oca; the major compounds
shaped with granular sizes between 20 and 55 μm, were malvidin 3-O-glucoside and malvidin
amylose content of 18.4 %, amylopectin with 3,5-O-diglucoside; the rest were petunidin
average chain length of 22.4 and β-amylolysis 3,5-O-diglucoside, peonidin 3,5-O-diglucoside,
limit of 64.5 %, B-type X-ray diffraction pattern, petunidin 3-O-glucoside, delphinidin 3-O-glucosides
gelatinisation enthalpy of 14.6 J/g and peak tem- and peonidin3-O-glucoside (Alcalde-Eon et al.
perature of the endothermic DSC (differential 2004).
scanning calorimetry) transition was 55.9 °C Caffeic, vanillic and cinnamic acid deriva-
(Santacruz et al. 2002). Santacruz et al. (2003) tives, flavan-3-ols and flavones derivatives, were
found that oca starch gel stored at 4 °C showed a identified as the main non-anthocyanin phenolic
high increase in G′ during the first day of storage. compounds in the aqueous tuber fraction of two
A decrease in pH from 6.5 to 4.0 produced a loss different coloured oca genotypes (Chirinos et al.
of structure in the starch gel, as was showed by 2009). Acid hydrolysis revealed the presence of
the reduction of G′. Storage at freezing tempera- vanillic, caffeic and cinnamic acids and malvidin
ture (−20 °C) produced higher changes in G′ than in the aqueous fraction. The ethyl acetate tuber
refrigeration conditions. fraction was composed mainly of caffeic and cin-
Oxalis tuberosa 143

namic acid derivatives as well as flavan-3-ols, but markedly decreased contents of starch and
flavones and flavanone derivatives. The flavan-3- organic acid. The chemical composition (range,
ols, flavones and flavanones detected in both per 100 g edible matter) reported for fresh and
fractions corresponded to bound forms of cate- sunned tubers was respectively as follows: dry
chin, luteolin and apigenin and naringenin, matter 10.4–14.3 g, 16.1–20 g; starch 5.2–9.2 g,
respectively. The aqueous tuber fractions were 2.2–6.5 g; soluble solids 6.2–92, 6.7–9.7 oBrix;
the major contributors to the ABTS antioxidant total sugars 2119–3635 mg, 2910–5870 mg;
capacity (77–82 %). sucrose 1208–1826 mg, 1901–3955 mg; fructose
Storage at 4 °C substantially extended the 477–1085 mg, 553–1217 mg; glucose 433–
storage life of oca tubers held for 20 weeks (Kays 1025 mg, 456–988 mg; organic acids 2567–
et al. 1979). Respiratory rate (4 mg CO2/kg/h at 3385 mg, 937–1970 mg; glutarate
4°), fresh weight and colour loss, and sprouting 1071–1578 mg,121–516 mg; malate 636–
were retarded. Composition changes of the tubers 1088 mg,139–660 mg; oxalate 306–539 mg,251–
(% reducing sugars, non-reducing sugars, total 451 mg; succinate 270–375 mg, 294–364 mg and
sugars, starches, total carbohydrates, total nitro- tartarate 57–138 mg, 56–191 mg. Twelve South
gen and non-protein nitrogen) were less pro- American and two New Zealand cultivars of oca
nounced at 4 °C than at 21 °C. Tubers stored at were found to contain total oxalate levels of
21 °C underwent substantial changes in composi- 80–194 mg/100 g wet matter (WM) in raw tubers
tion, the majority of which tended to coincide (Sangketkit et al. 2001). Oxalate levels ranged
with sprout formation. The mean dry matter con- from 77 to 220 mg/100 g WM for boiled and
tent of freshly harvested oca tubers was steamed tubers, while the levels of oxalate found
14.3 g/100 g fresh weight (FW), and after storage in baked tubers were significantly increased
for 6 weeks at 16.4 °C was 14.6 g/100 g FW when compared to the raw tubers. The oxalate
(Savage et al. 2008). The mean soluble oxalate content of the baked tubers ranged from 164 to
content of freshly harvested tubers was 335 mg/100 g WM. Total calcium content of the
162.1 mg/100 g FW, and, after 6 weeks storage, raw tubers ranged from 7.5 to 15.5 mg/100 g
173.5 mg/100 g FW. WM; cooking had a little effect on the calcium
Six organic acids had been reported in oca content. Among cultivars mean oxalate/calcium
tubers: ascorbic, malic, tartaric, succinic, glutaric ratio of the raw, boiled and steamed tubers ranged
and oxalic acids (Hermann and Erazo 2000; Ross from 2.5 to 9.9. Baked tubers had a significantly
et al. 1999; Bradbury and Emshwiller 2011). Of increased oxalate/calcium ratio (mean for all cul-
these oxalic acid is most deleterious to human tivars 9.5). Albihn and Savage (2001a, b) found
health (Ross et al. 1999; Albihn and Savage that oca tuber contained moderately high levels
2001a, b; Sangketkit et al. 2001). King (1988) of oxalate which existed in soluble form only.
reported low levels of 20.3–50.3 mg/100 g FW They found that oca tuber skin contained signifi-
oxalic acid in the tubers; in contrast, Ross et al. cantly higher oxalate level than the flesh 7.3 g/kg
(1999) reported levels of 92–221 mg/100 g FW in fresh weight (FW) compared to 1.7 and 1.4 g/kg
12 South American and two New Zealand culti- FW in the outer and inner flesh, respectively).
vars of oca. Levels of soluble and total oxalate The highest concentration was found in the skin
extracted from the tubers were not significantly of the pinkish-red cultivar (10.9 g/kg FW). All
different, suggesting that no calcium oxalate was cooking methods appeared to cause a migration
formed in the tuber. Hermann and Erazo (2000) of oxalate from the skin to the underlying flesh.
reported the highest oxalate value of 306– On a fresh weight basis, baking significantly
539 mg/100 g FW in freshly harvested tubers and increased the oxalate concentration in the whole
251–451 mg/100 g in sunned tubers. Sunning tuber, whereas boiling decreased the concentra-
reduced oxalate levels by an average of 26 %. In tion and steaming had no significant effect.
comparison to fresh tubers, sunning increased Boiling might therefore be a better way of cook-
dry matter content, soluble solids and total sugars ing oca than baking when a low intake of oxalate
144 Oxalidaceae

is desired. They found that the bioavailability of Antioxidant Activity

oxalate varied among individuals and depended
on other constituents of a combined meal. Oca For oca tubers, the hydrophilic antioxidant capac-
consumption with sour cream was found to sig- ity (HAC) and lipophilic antioxidant capacity
nificantly decrease the uptake of oxalate. On a (LAC) as determined by ABTS (2,2′-azinobis(3-
dry matter basis, the mean soluble oxalate of the ethylbenzothiazoline-6-sulfonic acid) ) ranged
raw oca cultivars was 935.5 mg/100 g DM rising from 1637 to 4771 μg TE/g and 69 to 320μgTE/g,
to 1364.8 mg/100 g DM when baked; in contrast, respectively (Campos et al. 2006). HAC expressed
the mean vitamin C content of the raw tubers was on a phenolic basis ranged from 1935 to 3614 mg
109.8 mg/100 g DM falling to a mean of TE/g chlorogenic acid equivalent. Oca tubers pre-
62.4 mg/100 g DM when the tubers were baked sented HAC higher than those observed for blue-
(Dubois et al. 2007). The skin and the tissue of all berries and blackberries. HAC in oca was related
the cultivars were a very similar orange–yellow to total phenolic and total anthocyanin content.
when cooked. Total phenolic content of oca tubers ranged from
Fresh oca tubers were found to contain per 0.71 to 1.32 mg/g with the purple coloured variet-
100 g edible portion 10.4–14.3 g dry matter, 5.2– ies having higher content and the yellow coloured
9.2 g starch, 6.2–9.2 oBrix (soluble solids), 2119– varieties lower phenolic content. Total carotenoid
3636 mg total sugars, 1208–1826 mg sucrose, content range from 2 to 25 μg/g in oca and mois-
477–1085 mg fructose, 433–1025 mg glucose, ture content 80–85 %. Total anthocyanin content
2567–3385 mg organic acids, 1071–1578 mg ranged from 0.14 to 1.3 mg/g; anthocyanin was
glutarate, 636–1088 mg malate, 306–539 mg concentrated in the tuber skin. There was no cor-
oxalate, 270–315 mg succinate and 57–138 mg relation between LAC and total carotenoid con-
tartarate (Hermann and Erazo 2000 ). Sun-dried tent in oca tubers, the lipophilic fraction
oca tubers were found to contain 16.1–20 g dry contributed 1.9–10.2 % to the total antioxidant
matter, 2.2–6.5 g starch, 6.7–9.7 oBrix (soluble capacity values for oca. According to the results
solids), 2910–5870 mg total sugars, 1901– obtained, the HAC range values for the crops
3955 mg sucrose, 553–1217 mg fructose, 456– studied followed the descending order
988 mg glucose, 937–1970 mg organic acids, mashua ≥ oca ≥ native potato ≥ ulluco.
121–516 mg glutarate, 139–660 mg malate, 251–
451 mg oxalate, 294–364 mg succinate and
56–191 mg tartarate. Sun-drying reduced oxalate Antimicrobial Activity
levels in dry matter by an average of 26 % and
also sharply reduced starch content, organic A protein of 18 kDa present in all oca morphot-
acids, glutarate and malate but significantly ypes was found to be the most abundant protein
increased total soluble solids, total sugars, in oca tubers (20–40 % of total soluble tuber pro-
sucrose, fructose and glucose. tein) (Flores et al. 1999, 2002). This protein, oca-
Hairy root cultures of O. tuberosa transformed tin, appeared to be a tuber-specific protein
with Agrobacterium rhizogenes exuded constitu- localized in the parenchyma cells of the pith and
tive levels of harmine (7-methoxy-1-methyl-β- the peridermis, representing a good source of
carboline) and harmaline (3,4-dihydroharmine), amino acids for the tuber. In addition to its stor-
the main fluorescent compounds detected from age role, ocatin was very active against the fol-
oca’s root exudates (Bais et al. 2003, 2002). lowing fungal pathogens: Rhizoctonia solani,
Transformed roots showed better growth and Phytophthora cinnamomi, Fusarium oxysporum
exudation of harmine and harmaline compared to and Nectria haematococcus. Anti-bacterial activ-
the untransformed normal roots. ity was also found against Agrobacterium tume-
Oxalis tuberosa 145

faciens, A. radiobacter, Serratia marcescens and model for studying domestication resulting in oppos-
ing crop phenotypes. Econ Bot 65(1):76–84
Pseudomonas aureofaciens, and the gene encod-
Campos D, Noratto G, Chirinos R, Arbizu C, Roca W,
ing ocatin was isolated and observed to have high Cisneros-Zevallos L (2006) Antioxidant capacity and
homology with pathogenesis-related (PR) pro- secondary metabolites in four species of Andean tuber
teins found in a wide variety of species (Flores crops: native potato (Solanum sp.), mashua
(Tropaeolum tuberosum Ruiz & Pavón), Oca (Oxalis
et al. 2002).
tuberosa Molina) and ulluco (Ullucus tuberosus
Caldas). J Sci Food Agric 86:1481–1488
Chirinos R, Betalleluz-Pallardel I, Huamán A, Arbizu C,
Other Uses Pedreschi R, Campos D (2009) HPLC-DAD charac-
terisation of phenolic compounds from Andean oca
(Oxalis tuberosa Mol.) tubers and their contribution to
Oca plants, tubers and foliage can also be used as the antioxidant capacity. Food Chem
stock feed for livestock such as pigs which relish 113(4):243–1251
both tubers and foliage. Oca is also grown as Cortella AR, Pochettino ML (1995) Comparative mor-
phology of starch of three Andean tubers. Starch/
home garden ornamentals.
Stärke 47:455–461
Dubois M, Savage GP, Martin RJ (2007) The effect of
cooking on the composition and colour of New
Comments Zealand grown oca. Food Chem 104(2):768–773
Duke JA (1994 onwards) Dr. Duke’s phytochemical and
ethnobotanical databases.
Oca is usually propagated vegetatively by plant- duke/plants.html
ing whole tubers and is a common component of Emshwiller E, Doyle JJ (2002) Origins of domestication
a crop rotation system with potato, atost or faba and polyploidy in oca (Oxalis tuberosa: Oxalidaceae).
2. Chloroplast-expressed glutamine synthetase data.
Am J Bot 89(7):1042–1056
Emshwiller E, Theim T, Grau A, Nina V, Terrazas F
(2009) Origins of domestication and polyploidy in oca
Selected References (Oxalis tuberosa; Oxalidaceae). 3. AFLP data of oca
and four wild, tuber-bearing taxa. Am J Bot
Albihn PBE, Savage GP (2001a) The bioavailability of
Flores HE, Vivanco JM, Loyola-Vargas VM (1999)
oxalate from oca (Oxalis tuberosa). J Urol
Radicle biochemistry: the biology of root-specific
metabolism. Trends Plant Sci 4:220–226
Albihn PBE, Savage GP (2001b) The effect of cooking on
Flores T, Alape-Girón A, Flores-Díaz M, Flores HE
the location and concentration of oxalate in three cul-
(2002) Ocatin. A novel tuber storage protein from the
tivars of New Zealand‐grown oca (Oxalis tuberosa
Andean tuber crop oca with antibacterial and antifun-
Mol). J Sci Food Agric 81(10):1027–1033
gal activities. Plant Physiol 128(4):1291–1302
Alcalde-Eon C, Saavedra G, Pascual-Teresa SD, Rivas-
Flores HE, Walker TS, Guimaraes RL, Bais HP, Vivanco
Gonzalo JC (2004) Liquid chromatography–mass
JM (2003) Andean root and tuber crops: underground
spectrometry identification of anthocyanins of isla oca
rainbows. Hortscience 38:161–167
(Oxalis tuberosa, Mol.) tubers. J Chromatogr A
Groen LE, Siemonsma JS, Jansen PCM (1996) Minor
species yielding non-seed carbohydrates. In: Flach M,
Arbizu C, Tapia M (1994) Andean tubers. In: Bermejo
Rumawas F (eds) Plant resources of South-East Asia
HJE, León J (eds) Neglected crops: 1492 from a dif-
No 9, Plants yielding non-seed carbohydrates. Prosea
ferent perspective, Plant production and protection
Foundation, Bogor, pp 165–186
services series no. 26. FAO, Rome
Gross R, Koch F, Malga I, Miranda AF, Schoeneberger H,
Bais HP, Park SW, Stermitz FR, Halligan KM, Vivanco
Trugo LD (1989) Chemical composition and protein
JM (2002) Exudation of fluorescent beta-carbolines
quality of some local Andean food sources. Food
from Oxalis tuberosa L roots. Phytochemistry
Chem 34:25–34
Hermann M, Erazo C (2000) Compositional changes of
Bais HP, Vepachedu R, Vivanco JM (2003) Root specific
oca tubers following postharvest exposure to sunlight.
elicitation and exudation of fluorescent β-carbolines in
CIP Program report 1999–2000, pp 391–396
transformed root cultures of Oxalis tuberosa. Plant
Hernández-Lauzardo AN, Méndez-Montealvo G,
Physiol Biochem 41(4):345–353
Velázquez del Valle MG, Solorza-Feria J, Bello-Pérez
Bradbury EJ, Emshwiller E (2011) The role of organic
LA (2004) Isolation and partial characterization of
acids in the domestication of Oxalis tuberosa: a new
146 Oxalidaceae

Mexican Oxalis tuberosa starch. Starch-Stärke Sangketkit C, Savage GP, Martin RJ, Mason SL (2001)
56(8):357–363 Oxalate content of raw and cooked oca (Oxalis
Kays SJ, Gaines TP, Kays WR (1979) Changes in the tuberosa). J Food Comp Anal 14(4):389–397
composition of the tuber crop Oxalis tuberosa Molina Santacruz S, Koch K, Svensson E, Ruales J, Eliasson AC
during storage. Sci Hortic 11(1):45–50 (2002) Three underutilised sources of starch from the
King SR (1988) Economic botany of the Andean tuber Andean region in Ecuador Part I. Physico-chemical
crop complex: Lepidium meyenii, Oxalis tuberosa, characterisation. Carbohydr Polym 49(1):63–70
Tropaeolum tuberosum and Ullucus tuberosus. Ph.D. Santacruz S, Ruales J, Eliasson AC (2003) Three under-
dissertation, City University of New York. (cited by utilised sources of starch from the Andean region in
Bradbury and Emshwiller 2011) Ecuador. Part II. Rheological characterisation.
King SR, Bastien HHC (1990) Oxalis tuberosa Mol. Carbohydr Polym 51(1):85–92
(Oxalidaceae) in Mexico: an Andean tuber crop in Savage GP, Mason SL, Vanhanen L (2008) The effect of
Meso-America. Adv Econ Bot 8:77–91 storage on the oxalate content of New Zealand
King SR, Gerhoff SN (1987) Nutritional evaluation of grown oca. Int J Food Sci Technol 43(12):
three under-exploited Andean tubers: Oxalis tuberosa 2130–2133
(Oxalidaceae), Ullucus tuberosus (Basellaceae) and Sperling CR, King SR (1990) Andean tuber crops: world-
Tropaeoleum tuberosum (Tropaeolaceae). Econ Bot wide potential. In: Janick J, Simon JE (eds) Advances
41(4):503–511 in new crops. Timber Press, Portland, pp 428–435
Martin RJ, Savage GP, Deo B, Halloy SRP, Fletcher PJ Stegemann H, Majino S, Scmiediche P (1988)
(2005) Development of new oca lines in New Zealand. Biochemical differentiation of clones of oca (Oxalis
Acta Hort (ISHS) 670:87–92 tuberosa) by their tuber proteins and the properties of
Popenoe H, King SR, Léon J, Kalinkowski LS, Vietmeyer these proteins. Econ Bot 42:37–44
ND, Dafforn M (1989) Lost crops of the Incas. Little The Plant List (2013) Version 1.1. Published on the
known plants of the Andes with promise for world- Internet;.
wide cultivation. National Academy Press, Trognitz BR, Hermann M (2001) Inheritance of tristyly in
Washington, DC Oxalis tuberosa (Oxalidaceae). Heredity (Edinb)
Ross AB, Savaga GP, Martin RJ, Vanhanen L (1999) 86(Pt 5):564–573
Oxalates in oca (New Zealand Yam) (Oxalis tuberosa Valcárcel-Yamani B, Rondán-Sanabria GG, Finardi-Filho
Mol.). J Agric Food Chem 47:5019–5502 F (2013) The physical, chemical and functional char-
Sangketkit C, Savage GP, Martin RJ, Searle BP, Mason acterization of starches from Andean tubers: Oca
SL (2000) Sensory evaluation of new lines of oca (Oxalis tuberosa Molina), olluco (Ullucus tuberosus
(Oxalis tuberosa) grown in New Zealand. Food Qual Caldas) and mashua (Tropaeolum tuberosum Ruiz &
Pref 11(3):189–199 Pavón). Braz J Pharm Sci 49(3):453–464
Piper methysticum

Scientific Name Estonian: Kavapipar

Fiji: Yangona, Yaqona
Piper methysticum G. Forster Finnish: Kavakava
French: Ava, Kava, Kava Kava, Kawa-Kawa
German: Kava-Kava, Kavapfeffer, Kawa-Kawa,
Synonyms Kawa-Pfeffer, Kawapfeffer, Rauschpfeffer
Hawaiian: ‘Awa
Methysticum methysticum (G.Forst.) A.Lyons Hungarian: Kava, Kávacserje, Mámorbors
Irian Jaya: Bari, Dikoi, Ikawati, Tigwa, Waghi,
Family Italian: Pepe Kava
Marquesan: Kava, Kav-Kava
Piperaceae Niuean: Kavainu
Papua New Guinea: Ayuw, Bari, Bikwe, Dikoi,
Gamada, Gamoda, Gumada, Irka, Jeliki, Ka,
Common/English Names Karangimi, Kau, Keu, Koi, Komata, Koniak,
Koriar, Kurar, Oyo, Sagainya, Sika, Tigwa,
Awa, Kava, Kava-Kava, Kava Pepper, Kava Toa, Toe, Towe, Tue, Tui, Tokarabu, Uati,
Shrub, Kawa, Yangona Pepper Waghi, Wariki, Wati
Pohnpei: Sakau
Polish: Pieprz Kawakawa; Pieprz Metystynowy
Vernacular Names Samoa: ‘Ava, Ava Ava, Kava
Slovašcina: Kavakava
Arabic: Fulful Kâwah Spanish: Kava, Kava Kava, Kavaka, Kawa
Brazil: Cava Cava Swedish: Kava Kava
Chinese: Ka Wa Hu Jiao Tahiti: Ava, Ava-Ava, Evava
Czech: Pepřovník Opojný Tongan: Kava, Kav-Kava
Danish: Kava, Kava Kava, Kavarod Tubuai: Ava
Esperanto: Kavao Turkish: Kava Biberi, Kawa Kawa

© Springer International Publishing Switzerland 2016 147

T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_15
148 Piperaceae

Vanuatu: M Nigui (Hiu), Maloku (Maewo), Agroecology

Namaloku (Nguna), Sini (Central Pentecost)
Mele (south Pentecost), Maloku (North Piper methysticum being a tropical species
Pentecost), Nga (Ureparapara), Naga (Mota requires tropical conditions to flourish. It thrives
Lava), Gwie (Vanua Lava), Malop (Mere best in loose, well-drained, moist, well-
Lava), Amaloku (West Ambae), Maloku (East composted soils in a protected, partially shaded
Ambae), Bir (Central Santo), Naxai (North position, between 300 and 500 m above sea level
Santo), Malou (Southwest Santo), Malohu in a tropical environment of 20–35 °C, 70–100 %
(West Santo), Hae (Malo), Malox (North relative humidity and rainfall above 2000 mm
Malakula), Nem Leu, Melo, Melu, Malok per annuum.
(Northeast Malakula), Malk, Malox
(Northwest Malakula), Nimvulm (Southwest
Malakula), Maix, Namonggomongg, Edible Plant Parts and Uses
Monggmongg, Merox (Southeast Malakula),
Meruh (East Malakula), Lewewe Ndrame The root and rhizome (underground stem) of
(North Ambryn) Vatimeai (North Panama) kava are used to prepare beverages, extracts, cap-
Malou (South Panama), Malk, Miau, Mia, sules, tablets, and topical solutions (NCAM
Mio, Mak (West Epi), Namaluk (Tongoa), 2006). Its thick roots and stumps (stem bases) are
Namaloku (Emae) Malok (Makira), Namloku shaved, mashed or grounded into powder
(Nguna) (Plates 2 and 3) and made into a cold beverage
Wallis and Futuna: Kava used similarly to alcohol and has a sedative
effect. Kava is widely and commonly consumed
as a social beverage to establish kinship in the
Pacific island communities (McDonald and
Origin/Distribution Jowitt 2000). The roots or shavings can also be
chewed. Chewing produces the strongest effect
The species is indigenous to and cultivated in the because it produces the finest particles.
islands of south Pacific, from Hawaii to Papua
New Guinea, with the notable exception of New
Caledonia, New Zealand and most of the Solomon Botany
Islands (Singh 1992).
Phytochemical studies conducted by Lebot An evergreen, robust, dioecious, vine-like peren-
and Lèvesque (1989) to elucidate the origin of nial shrub, 1–4 m high, with glabrous, woody,
this Oceania plant found the lineage of chemical branching stems and massive base with lateral
chemotypes suggested that Piper wichmanii was roots to 2 m long and 8 cm diameter. Leaves
the wild species from which farmers domesti- green, alternate, glabrous, cordate, 10–30 cm by
cated cultivars of Piper methysticum. Based on 9–24 cm, entire, apex acute, palmately veined
morphological, chemical, cytological and genetic and borne on 2–7 cm long petioles (Plate 1). The
evidence demonstrating the absence of taxo- flowers are small, unisexual, whitish, occurring
nomic distinction between Piper methysticum in short, irregular, axillary spikes or opposite
and Piper wichmannii, Lebot and Levesque leaves. The floral bracts are rounded and peltate
(1996) concluded that Piper methysticum is not a on pubescent pedicels. The male spikes with
separate species, but rather a group of sterile cul- flowers bearing two short stamens, female spikes
tivars selected from somatic mutants of P. wich- (rare) female flower with a unilocular ovary and
mannii. As P. methysticum was described first stigma. The fruit, a berry is seldom produced and
(1786), it had priority making P. wichmannii the plant must be propagated by dividing the
(1910) superfluous. roots or through stem cuttings.
Piper methysticum 149

Nutritive/Medicinal Properties Mg 0.099 %, Na 0.111 %, Fe 0.017 %, Zn

22.07 ppm, Mn 7.30 ppm, Cu 15 ppm, cl
Phytonutrients 0.012 ppm, S 0.018 ppm, B 11.6 ppm (Lebot and
Cabalion 1988).
Fresh kava root was reported to contain 43 %
starch, 20 % fiber, 12 % water, 3.2 % sugars,
3.6 % proteins, 3.2 % minerals and up to 20 % Phytochemicals – Kavalactones/
kava lactones (Leung and Foster 1996). The com- Chalcones
position of kava drug from kava basal stem and
roots in Fiji was determined to have respectively: More than 40 compounds belonging to the classes
moisture 14.60, 13.66%; carbohydrates 74.11, of kavapyrones, alkaloids, steroids, chalcones,
64.22 %; fibre 10.66, 13.09 %; ash 2.33, 3.57 %; long-chained fatty acids and alcohols have been
K 2.161, 2.00 %; Ca 0.455, 0.201 %; mg 0.105, isolated and identified from Piper methysticum
0.179 %; Na 0.0362, 0.060 %; Al 0.0202, 0.15 %; (Parmar et al. 1997). Van Veen (1939) isolated a
Fe 0.017, 0.106 %; Zn 60.66, 55.35 ppm; Mn
12.04, 51.77 ppm; Cu 13,08, 16.95 ppm (Duve
and Prasad 1983). Dry commercial kava powder
was analysed by IRCC (Institut de Rechrches sur
le Café et le Cacao) laboratory in Vanuatu to have
sugars 0.50 % sucrose, 0.10 % maltose, 1.75 %
fructose and 0.85 % glucose; amino acids –
aspartic acid 0.28 %, threonin 0.08 %, serine
0.11 %, glutamic acid 0.26 %, glycine 0.11 %,
phenylalanine 0.07 %, histidine 0.05 %, lysine
0.10 %, arginine 0.08 %, alanine 0.16 %, valine
0.11 %, methionine 0.02 %, isoleucine 0.07 %,
leucine 0.14 % and tyrosine 0.06 %; kavalactones
5.23 %; dihydro-5,6 kavalactones plus dehydro-
5,6-kavalctones; minerals K 2.37 %, Ca 0.372 %, Plate 1 Cordate leaves of young kava seedling

Plate 2 Dried kava roots (right) and shavings (left)

150 Piperaceae

Plate 3 Pure pounded kava powder

narcotic substance from dried kava stems and The content (weight %) of six major kavalac-
roots, and called it marindinin which later was tones in the dried kava roots was reported as:
found to be identical with dihydrokawain of kavain 2.58 %, dihydrokavain 1.37 %, methysti-
Borsche. The concentrations of desmethoxyyan- cin 1.82 %, dihydromethysticin 1.89 %, yangonin
gonin, yangonin, dihydromethysticin, methysticin, 1.73 %, desmethoxyyangonin 0.81 % analysed
dihydrokawain and kawain in kava ether extract in Hawaii (Young et al. 1966); kavain 1.90 %,
were determined (Young et al. 1966). Dutta et al. dihydrokavain 2.37 %, methysticin 2.12 %, dihy-
(1972) isolated yangonin and methysticin from dromethysticin 1.12 %, yangonin 1.21 %, desme-
kava roots. Recoveries of known compounds were thoxyyangonin 0.59 % analysed in Fiji (Duve
80–85 % for yangonin and 90–95 % for all others. 1981); kavain 2.30 %, dihydrokavain 3.28 %,
Fresh kava root stock contained 80 % water, while methysticin 2.06 %, dihydromethysticin 1.60 %,
dried rootstock consists of approximately 43 % yangonin 1.16 %, desmethoxyyangonin 0.91 %
starch, 20 % fibre, 12 % water, 3.2 % sugars, 3.6 % analysed in Vanuatu (Lebot and Lévesque 1996).
proteins, 3.2 % minerals and 15 % (3–20 %) kava- The composition of the major kavalatones in kava
lactones (Lebot et al. 1992). Kavalactone content root, stem and leaf were reported respectively by
was reported to be greatest in the roots and to Smith (1983) askavain (34.5, 0.6, 2.5 %); dihy-
decrease higher up the plant. Relative concentra- drokavain (17.1,23.2,69.8 %); methysticin (20.8,
tions of 15 %, 10 % and 5 % have been observed 13.9, 0.8 %); dihydromethysticin (5.3, 59.6,
in the root, stumpand basal stems, respectively. 22.5 %); yangonin (0.8,0.8, 1.2 %) and desme-
Kavalactones (amounting to %) had been reported thoxyyangonin (21.6, 1.8, 3.0 %).
in all parts of the kava plant comprising the major The first compound isolated from Piper
ones: kavain (1.8 %), methysticin (= kavahine, methysticum was methysticin (Cuzent 1981) also
kavakin, kavatin, kanakin) (1.2 %), demethoxy- known as kavatin, kavahin and kanakin. The
yangonin (1 %), yangonin (1 %), dihydrokavain IUPAC has recommended the name 5,6-dihydro-
(0.6 %), dihydromethysticin (0.5 %) and traces of 4-methoxy-6-[3′,4′-(methylenedioxy)styryl]-2H-
11,12-dimethoxyhydrokavain, 11-hydroxy12- pyran-2-one but chemical abstracts indexing has
methoxykavain, 11-methoxy-nor-yangonin, adopted the latter path, using 5-hydroxy-3-
11-methoxy-yangonin and the two ethylketones methoxy-7-[3,4-methylenedioxyphenyl]-2,6-
cinnamoylacetone and methylendioxy-3,4- hepta-dienoic acid- lactone. The ethylenic
cinnamoylidenacetone (Shulgin 1973; Young et al. dihydroanalog dihydromethysticin was isolated
1966). almost 50 years later by Winzheimer (1908).
Piper methysticum 151

Nolting and Kopp (1874) isolated yangonin. The A total of 19 kavalactones namely 10-
most extensive studies on the constituents of kava methoxyyangonin; hydroxykavain; dihydro-
plant were conducted by Borsche and coworkers 5,6-dehydrokavain; 7,8-dihydrokavain; 7,8-
in 13 papers wherein they reported the isolation dihydroyangonin; kavain; 7,8-dihydro-5-
and structure of two compounds kawain (also hydroxykavain; 5,6-dihydroyangonin; 11-
known as kavain and gonosan) and dihydro- h y d r oxy-12-methoxydihydrokavain;
kawain and the structural elucidation of the crys- 11-methoxyyangonin; 5,6-dehydrokavain (des-
talline components methysticin, dihydromethysticin methoxyyangonin); 5,6,7,8-tetrahydroyango-
and yangonin (Borsche 1927; Borsche and nin; methysticin; dihydromethysticin; 11,12
Bodenstein 1929; Borsch and Blount 1930, 1932, dimethoxydihydrokavain; yangonin; 11-methoxy-
1933; Borsche and Gerhard 1914; Borsch and 12-hydroxydehydrokavain; 11-hydroxyyango-
Lewinsohn 1933; Borsche et al. 1927a, b, 1929a, nin; 5,6-dehydromethysticin were identified in
b; Borsche and Peitzsch 1929a, b, 1930; Borsche kava roots (Hänsel 1968; Lebot and Cabalion
and Roth 1921; Borsche and Walter 1927). 1988; He et al. 1997; Whittaker et al. 2008;
Yangonin (= 4-methoxy-6-(p-methoxy-β-styryl)- Dharmaratne et al. 2002; Xuan et al. 2008;
α-pyrone) was not so recognised by Borsche and Teschke 2011; WHO 2007), with flavokavains A,
co-workers (Borsche and Gerhard 1914; Borsche B and C (He et al. 1997); kava lactones (Hänsel
and Bodenstein 1929). They had postulated it as et al. 1968) identified in kava roots included:
a γ-pyrone, although they had correctly inter- kavain; 7,8-dihydrokavain; 5,6-dehydrokavain;
preted the structures of all other substances iso- yangonin; 5,6,7,8-tetrahydroyangonin; methysti-
lated from kava (Borsche et al. 1927a, b; Borsche cin; dihydromethysticin; 5,6-dehydromethysticin;
and Peitzsch 1929a, 1930). Yangonin served as 5,6-dihydroyangonin; 7,8-dihydroyangonin; 10-
the basis for two additional substances the meta- methoxy-yangonin; 11-methoxy-yangonin; 11-
methoxy analogue 11-methoxyyangonin (Hänsel hydroxy-yangonin; hydroxykavain; 11-methoxy-
and Klaproth 1966) and the phenolic counterpart 12-hydroxy-dehydrokavain (Lebot and Cabalion
11-methoxy-nor-yangonin (Hänsel et al. 1966c) 1988). Major compounds identified in Kava
isolated from Kava-kava, both of which had been extracts by low-resolution GC/MS included
established synthetically as possessing the kava lactones like kavain, dihydro-and dehydro-
α-pyrone ring. The completely conjugated ana- kavains, yangonin and tetrahydroyangonin,
logue, 5,6-dehydromethysticin, was identifed as methysticin and dihydromethysticin (Nerurkar
a component of kava-kava by Hänsel (1968). The et al. 2004; Lopez-Avila and Benedicto 1997).
following kavalactones were isolated from kava Other kavalactones were identified as minor
roots: 5,6,7,8-tetrahydro-yangonin (Achenbach components including 1-methoxyyangonin
et al. 1971); 11-hydroxy-12-methoxy-dihydro- (Hänsel and Klaproth 1966), 11-metho
kawain and 11,12-dimethoxy-dihydrokawain xynoryangonin and 5,6-dehydromethysticin
(Achenbach et al. 1972). Two yellow pigment (Hänsel et al. 1966b); 1-methoxyyangonin
materials, flavokawin A and B, were isolated (Hänsel and Klaproth 1966), 11-methoxynoryan-
from kava rhizomes (Hänsel et al. 1961). The gonin and 5,6-dehydromethysticin (Hänsel et al.
structures of these were established by synthesis 1966b); 10-methoxyyangonin, 11-hydroxyyan-
to be substituted chalcones bearing an obvious gonin and 11-methoxy-12-hydroxydehydroka-
biogenetic relationship to the styrylpyrones vain (Duve 1981; He et al. 1997). Additionally,
(Hänsel et al. 1963). Hänsel and Schulz (1973) three hydro-derivatives of yangonin were identi-
synthesized 5,6,-cis-kavain 5-ol and 5,6-trans- fied from kava plant. These include 5,6-dihydroy-
kavain-5-ol. Flavokawin A was elucidated as angonin (Hänsel 1968; Duve 1981),
2′-hydroxy-4-4′-6′-trimethoxychalcone and fla- 7,8-dihydroyangonin (Duve 1981) and 5,6,7,8-tet-
vokawin B as 2′-hydroxy-4′-6′ dimethoxychal- rahydroyangonin (Achenbach et al. 1971; Hänsel
cone. The flavokavins A, B and C were reported 1968). Hydroxylated α-pyrone derivatives, includ-
to have a different skeleton (Duve 1976; Hänsel ing cis-5-hydroxykavain (Duve 1981) and
et al. 1963; Dutta et al. 1973, 1976). 5-hydroxydihydrokavain (Achenbach and
152 Piperaceae

Wittman 1970), had also been isolated from P. and 39 months. After 39 months of storage 93.9 %
methysticum as minor components. of dihydrokavain, 81.6 % of kavain, 72.4 % of
The kavalactone, 11-methoxy-5,6-dihydro- dehydrokavain, 54.9 % of tetrahydroyangonin,
yangoninand eight previously reported analogs 25.8 % of dihydromethysticin, 32.1 % of yangonin
along with four other aromatic compounds were and 29.5 % of methysticin had deteriorated in the
isolated from kava root extracts (Ranjith et al. powdered root samples.
2002). Nine kava lactones were detected from Concentrations of total kava lactones
kava roots using GC-MS including desmethoxyy- (methysticin, dihydromethysticin, kawain, dihy-
agonin, kavain, 7,8-dihydrokavain, hydroxyka- drokawain, yangonin and desmethoxyyango-
vain, yagonin, 5,6,7,8-tetrahydroxyyagonin, nin), ranged between 135 and 0.035 mg per
methysticin, dihydromethysticin and 11-hydroxy- serving in 10 beverage products, two chocolate
12-methoxydihydrokavain (Xuan et al. 2006). products, three unbrewed tea products, a drink
Quantities of desmethoxyyagonin, kavain, mix product tested and between 40 and 61 mg
7,8-dihydrokavain, yagonin, methysticin and per serving for the three dietary supplement
dihydromethysticin detected were 4.3, 6.9, products tested using LC-UV and LC-MS anal-
18.6, 5.7, 1.4 and 5.4 mg/g of dry weight, respec- yses (de Jager et al. 2004). One percent of kava
tively. Eighteen kava lactones, cinnamic acid bor- extract could be detected in food supplements
nyl ester and 5,7-dimethoxy-flavanone, and seven using liquid chromatography-atmospheric pres-
compounds, including 2,5,8-trimethyl-1-naphthol; sure chemical ionisation tandem mass spec-
5-methyl-1-phenylhexen-3-yn-5-ol; 8,11-octa- trometry, corresponding to approximately
decadienoic acid-methyl ester; 5,7-(OH)(2)-4′- 0.05–0.2 mg/g of the individual kava lactones
one-6,8-dimethylflavanone; pinostrobin chalcone; kavain, dihydrokavain, yangonin, desmethoxyy-
and 7-dimethoxyflavanone-5-hydroxy-4′, were angonin, methysticin and dihydromethysticin
identified from kava roots (Xuan et al. 2008). (Bobeldijk et al. 2005). Reliable quantification
Glutathione (26.3 mg/g) was found in the water was obtained from concentrations of
extract. Dihydro-5,6-dehydrokavain (DDK) was 0.25–1 mg/g, depending on the compound.
present at a higher level than methysticin and des- Reliable quantification was obtained from con-
methoxyyagonin, indicating that DDK to be a centrations of 0.25–1 mg/g, depending on the
major constituent of kava roots. compound. Gautz et al. (2006) found the combi-
Of 18 kavalactones or kava pyrones found in nation of near-infrared reflectance spectroscopy
kava, kawain, dihydrokawain, methysticin, and partial least-squares methods to be a conve-
dihydromethysticin, yangonin and desmethoxyy- nient, versatile and rapid analytical tool for
angonin were the six major kavalactones determination of kavalactones in dried kava
(Whittaker et al. 2008). The following kavalactone powder.
compounds were isolated from kava roots: Major and trace constituents of kava resin
kavalactones:10-methoxyyagonin; hydroxykavain; were identified using methane negative ion
dihydro-5,6-dehydrokavain; 7,8-dihydrokavain; chemical ionization (NICI) mass spectrometry
7,8-dihydroyagonin; kavain; 7,8-dihydro-5- (Duffield and Lidgard 1986; Duffield et al. 1986).
hydroxykavain; 5,6-dihydroyagonin; 11-hydroxy- Several of the kava lactones (kawain, methysti-
12-methoxydihydrokavain; 11-methoxyyagonin; cin, desmethoxyyangonin and yangonin) pro-
desmethoxyyagonin, 5,6,7,8-tetrahydroyagonin; duced abundant molecular anions when analysed
methysticin; dihydromethysticin; 11,12-dimethy- by methane NICI mass spectrometry, in contrast
oxydihydrokavain; yagonin; 11-methoxy-12-hy- to their 7,8-dihydro analogues which ionized
droxydehydrokavain; 11-hydroxyyagonin; and with poor efficiency to yield weak [M − H]−
5,6-dehydromethysticin (Xuan et al. 2008). Duve anions. Several new trace compounds formally
and Prasad (1983) investigated the stability of kav- resulting from decarboxylation of the 4-methoxy-
alactones in powdered kava root stored in screw- 2-pyrone ring system in the known compounds
capped glass bottles at room temperature for 22, 36 were also found. A simultaneous HPLC separa-
Piper methysticum 153

tion of the enantiomers of kavain, dihydrokavain, For ground kava (250–500 μm), 2 h of sub-
methysticin and dihydromethysticin as well as critical water extraction was required for a com-
the achiral dienolides yangonin and desme- plete extraction of kava lactones at 100 °C, while
thoxyyangonin was carried out on a ChiraSpher at 175 °C, 20 min were sufficient (Kubátová et al.
NT column (Boonen et al. 1997; Häberlein et al. 2001). For a complete extraction of the unground
1997). Using reverse-phase high-performance (shredded) kava, the time of extraction was
liquid chromatographic method, six major kava- extended to 40 min at 175 °C. Boiling for 2 h and
lactones [methysticin, dihydromethysticin extraction with Soxhlet apparatus for 6 h, both of
(DHM), kavain, dihydrokavain (DHK), deme- which employed water at atmospheric pressure,
thoxyyangonin (DMY) and yangonin] in Piper produced yields 40–60 % lower than those
methysticum (kava) were simultaneously deter- obtained with subcritical water. With unground
mined (Shao et al. 1998). The detection limits kava, 40 min of subcritical water extraction
(S/N = 3) for methysticin, DHM, kavain, DHK, yielded essentially the same recoveries of lac-
DMY and yangonin are approximately 0.5, 1.1, tones as 18 h of sonication with acetone, methy-
0.7, 1.1, 0.6 and 0.6 μg/ml, respectively. The lene chloride or methanol. Kava root extracted in
average recoveries are 100.2 % for methysticin, acetone yielded 100 % kavalactones, 96 % etha-
100.6 % for DHM, 100.0 % for kavain, 100.3 % nol yielded 100 % kavalactones, 25 % ethanol
for DHK, 98.9 for DMY and 98.2 % for yango- yielded 15 % kavalactones and water yielded
nin. A rapid micellar electrokinetic chromatogra- 2.97 % kavalactones (Denham et al. 2002).
phy method with diode-array detection was used Extraction rates also varied depending on the
to separate kavalactones kavain, dihydrokavain, temperature at which the products were prepared.
methysticin, dihydromethysticin, yangonin and Analysis of flavokavins in extracts of 172 sam-
demethoxyyangonin from dry kava extracts ples originating from four cultivars groups
(Lechtenberg Lechtenberg et al. 1999). (noble, medicinal, 2-days and wichmannii) indi-
Three kava lactone constituents of Piper cated that the ratio flavokavin B /kavalactones
methysticum, namely, kawain, methysticin and was much higher in 2 days (0.39) and wichman-
desmethoxyyangonin, were separated and identi- nii (0.32) compared to nobles (0.09) and medici-
fied by reverse-phase HPLC using superheated nal cultivars (0.10) (Lebot et al. 2014). For each
deuterium oxide as the mobile phase and on-line group, the ratios of flavokavins/kavalactones did
1H-NMR detection (Chienthavorn et al. 2005). not change significantly between roots, stumps or
Meissner and Häberlein (2005) used simultane- basal stems and among clones, indicating that
ous HPLC for the separation of six major kavapy- they were genetically controlled.
rones and the flavokavins A–C in an ethanolic
extract of Piper methysticum. Flavokavins A–C
contents of 0.62 mg/100 mg, 0.34 mg/100 mg Phytochemicals – Alkaloids/
and 0.14 mg/100 mg ethanolic kava extract was Flavonoids/Miscellaneous
determined respectively. Compounds identified Compounds
in commercial kava extracts by gas chromatogra-
phy (GC) and quadrupole time-of-flight mass There had been reports of isolation of alkaloids,
spectrometry (QTOFMS) with an electron flavonoids – flavokavins, sterols, organic acids,
ionization source: 7,8 dihyrokavain; kavain; aliphatic alcohols and other compounds from
5,6-dehydrokavain; 5,67,8-tetrahydroyangonin; kava plant. Three flavokavins (also known as fla-
7,8-dihydromethysticin; yangonin; methysti- vokawins or flavokavains), designated A, B and
cin; bornyl cinnamate; dihydroxy- C, had been identified from kava root (Dutta and
methoxyphenyl-phenyl-propen-one; hydroxyl- Ray 1973; Dutta et al. 1973, 1976; Hänsel et al.
dimethoxyphenyl-3-phenyl-2-propen-1-one; and 1961, 1963). Structures of these compounds had
hydroxyl-dimethoxyphenyl-3-methoxyphenyl-2- been confirmed by synthesis (Hänsel et al. 1963;
propen-1-one (Lopez-Avila and Yefchak 2009). Dutta et al. 1976; Dutta and Som 1978). Two
154 Piperaceae

alkaloids were isolated from a methanolic 1-ol, hexacosan-1-ol, hexadecane-1-ol, octadeca-

extract of kava root and were identified as nol-1ol and n-triacontane (Gracza and Ruff
1-cinnamoylpyrrolidine and 1-(methoxycinnam- 1986).
oyl)pyrrolidine (Achenbach and Karl 1970) and The root was also known to contain flavoka-
an alcohol dihydrokawain-5-ol (Achenbach and vins A and B, pipermethystine, cepharadione A,
Wittman 1970). A third alkaloid, pipermethys- cinnamalketone, and methylene dioxy-3,
tine, was isolated from leaves by Smith (1979; 4-cinnamalketone (Lebot et al. 1992). Bornyl
1983) and was found in small amounts in the cinnamate and a series of hydroxylated com-
stems and roots (Singh 1992). A few piperidine pounds resulting from the decarbonylation of the
alkaloids (i.e., pipermethystine, 3,4-epoxy-5- lactones had also been identified in kava resin by
pipermethystine, and awaine) were identified in methane chemical ionization GC/MS and two
the aerial parts of kava plant (stem peelings and previously described N-cinnamoyl pyrrolidine
leaves) (Dragull et al. 2003). Pipermethystine alkaloids along with stigmasterol (Cheng et al.
was concentrated in the stem peelings and leaves. 1988). Three dihydrochalcones: flavokavain A,
3,4-epoxy-5-pipermethystine and awaine were flavokavain B, flavokavain C; cinnamic acid bor-
new alkaloids with 3,4-epoxy-5-pipermethys- nyl ester and 5,7-dimethoxy-flavanone; and
tine found only in cv. Isa among the 11 cultivars compounds detected for the first time in kava
examined, and awaine occurred primarily in roots: 2,5,8-trimethyl-1-naphthol; 5-methyl-1-
young leaves of all cultivars. None of the three phenylhexen-3-yn-5-ol; 8,11-octadecadienoic
piperidine alkaloids were detected in the com- acid-methyl ester; 5,7-(OH)(2)-4′-one-6,8-
mercial root powders from Fiji, Tonga or dimethylflavanone; pinostrobin chalcone; and
Hawaii. Two conjugated diene ketones, cin- 7-dimethoxyflavanone-5-hydroxy-4′. Glutathione
namylidineacetone and 3,4-methylenedioxy- (26.3 mg/g) was found in the water extract (Xuan
cinnamylidineacetone (Jössang and Holho 1967), et al. 2008). Acetone was the most effective sol-
and organic acids (Achenbach and Karl 1971) vent in terms of maximum yield and types of
were isolated from kava roots. Kava plant had kava lactones isolated, followed by water and
been found to contain amides (2-methoxy cin- chloroform, whereas hexane, methanol and etha-
namic pyrrolide, cinnamic acid pyrollide), chal- nol were less effective as solvents. Total phenolic
cones (flavokavin A and B) and free and and antioxidant activity varied among the extract-
aromatic acids, anisic acid, benzoic acid, capri- ing solvents, with acetone and chloroform pro-
onic acid, hydroxyl cinnamic acid and deriva- ducing the highest effects, followed by water,
tives (Klohs 1967; Halzl et al. 1993). The kava while methanol, ethanol and hexane were less
methanol root extract yielded bornyl esters of effective. Two alkaloids, (−)-pipermethystine and
3,4-methylenedioxy cinnamic acid and cin- (−)-epoxypipermethystine, and two lactones,
namic acid, pinostrobin, flavokawain B and (+)-dihydromethysticin and yangonin, were iso-
5,7-dimethoxyflavanone (Wu et al. 2002a). Two lated from the kava plant (Naumov et al. 2008).
C-glycoside flavonoid compounds identified as
2″-O-rhamnosylvitexin and schaftoside were iso-
lated from kava leaves (Jhoo et al. 2007) and phy- Pharmacological Activities
tosterols such as stigmasterol, stigmastanol,
β-sitoserol and campesterol (Jössang and Molho Kava kavalactones were reported to possess phar-
1970; Gracza and Ruff 1986). Trace amount of a macological effects which include sedative, anx-
nitrogen-containing compound (cepharadione) iolytic, anti-stress, analgesic, local anaesthetic,
had also been isolated from the kava rootstock anticonvulsant and neuroprotective properties
(Jaggy and Achenbach 1992). Alipahtic and (Gounder 2006). Kava’s biological effects, due to
alicyclic alcohols had been isolated from the a mixture of compounds called kavalactones, had
rhizome: docosan-1-ol, docecanol-1-ol, eicosanol- been reported to include sedative, anxiolytic,
Piper methysticum 155

antistress, analgesic, local anaesthetic, anticon- Anticancer Activity

vulsant and neuroprotective properties (Singh
and Singh 2002). The pharmacological proper- Findings of a cancer incidence survey for the
ties of kava were postulated to include blockade Pacific Islands completed in the 1980s indicated
of voltage-gated sodium ion channels, enhanced that the more kava consumed by a population the
ligand binding to gamma-aminobutyric acid lower the cancer incidence for that population
(GABA) type A receptors, diminished excitatory (Steiner 2000). Crude kava extracts (dichloro-
neurotransmitter release due to calcium ion chan- methane and hexane fractions) showed good
nel blockade, reduced neuronal reuptake of nor- activity against A2780 ovarian tumour and K562
adrenaline (norepinephrine), reversible inhibition human leukaemia cancer cell lines in-vitro
of monoamine oxidase B and suppression of the (Tabudravu and Jaspars 2005). Bioassay-guided
synthesis of the eicosanoid thromboxane A(2), isolation afforded six known kava lactones dihy-
that was antagonistic to GABA(A) receptor dromethysticin; 7,8 dihydrokavain; kavain;
function. demethoxyyangonin; cis-yangonin and trans-
yangonin and two flavokavains A and B. The
anticancer of the fractions and eight compounds
Antioxidant Activity gave IC50 values ranging from 0.42 to 9.15 μg/ml
against K562 and 0.43–5.15 μg/ml against
The lipid oxidation assay did not reveal A2780.
antioxidant activities for kava root compounds Flavokawain A, B and C naturally occurring
demethoxyangonin, dihydrokawain, kawain, chalcones isolated from kava may hold promis-
dihydromethysticin or methysticin at 50 μg/ml ing anti-cancer effects as numerous studies
(Wu et al. 2002b). The antioxidant activities of revealed that both flavokawain A and B were
flavokawain A and yangonin could not be tested involved in the induction of cell cycle arrest in
in the lipid oxidation assay due to solubility prob- several cancer cell lines (Abu et al. 2013).
lems. However, yangonin and methysticin Flavokawain B was shown to be more effective in
showed moderate antioxidant activities in the treating in-vitro cancer cell lines as compared to
free radical scavenging assay at 2.5 mg/ml. flavokawain A and C. Flavokawain B also exerted
antinociceptive effects as well as anti-
inflammation properties. Flavokawain A, a chal-
Cyclooxygenase Enzyme Inhibitory cone from kava extract, induced apoptosis in
Activity bladder cancer cells by involvement of Bax
protein-dependent and mitochondria-dependent
Six compounds isolated from the ethyl acetate apoptotic pathway and suppresses bladder tumor
kava root extract were found to possess cyclo- growth in mice (Zi and Simoneau 2005). Studies
oxygenase inhibitory activity (Wu et al. 2002b). by Tang et al. (2008) found that kava flavokawain
Dihydrokawain and yangonin showed the high- A exerted different effects in bladder cancer cells
est COX-I and COX-II inhibitory activities at with wild-type versus mutant p53. In a p53 wild-
100 μg/ml, respectively. All kava compounds type, low-grade, and papillary bladder cancer cell
bornyl esters of 3,4-methylenedioxy cinnamic line (RT4), flavokawain A increased p21/WAF1
acid and cinnamic acid, pinostrobin, flavoka- and p27/KIP1, which resulted in a decrease in
wain B and 5,7-dimethoxyflavanone tested gave cyclin-dependent kinase-2 (CDK2) kinase activ-
good COX-I and moderate COX-II enzyme ity and subsequent G(1) arrest. In contrast, flavo-
inhibitory activities at 100 μg/mL (Wu et al. kawain A induced a G(2)-M arrest in six p53
2002a). Flavokawain B showed the highest mutant-type, high-grade bladder cancer cell lines
COX-I inhibitory activity at 100 μg/mL. (T24, UMUC3, TCCSUP, 5637, HT1376 and
HT1197). Flavokawain A significantly reduced
the expression of CDK1-inhibitory kinases, Myt1
156 Piperaceae

and Wee1, and caused cyclin B1 protein accumu- with minimum toxicity. Other kava constituents,
lation leading to CDK1 activation in T24 cells. including four kavalactones of similar structures
Suppression of p53 expression by small interfer- to methysticin, demonstrated minimum activities
ing RNA in RT4 cells restored Cdc25C expres- in inhibiting NF-kappaB. A potent chalcone ana-
sion and down-regulated p21/WAF1 expression, log, (E)-3-(3-hydroxy-4-methoxyphenyl)-1-
which allowed Cdc25C and CDK1 activation, (3,4,5-trimethoxyphenyl)prop-2-en-1-one
which then led to a G(2)-M arrest and an enhanced derived from flavokawain compounds found in
growth-inhibitory effect by flavokawain kava, dose-dependently inhibited A549 lung can-
A. Consistently, flavokawain A also caused a pro- cer cell viability, NF-κB, activation of caspases
nounced CDK1 activation and G(2)-M arrest in and activation of mitogen-activated protein
p53 knockout but not in p53 wild-type HCT116 kinases such as extracellular signal regulated
cells. kinase 1/2 (ERK1/2) and c-Jun N-terminal kinase
Studies showed that a 30-week kava treatment (JNK) (Warmka et al. 2012)
inhibited 4-(methylnitrosamino)-1-(3-pyridyl)- Studies by Tang et al. (2010) demonstrated
1-butanone plus benzo[a]pyrene-induced lung that flavokawain B (FKB), a kava chalcone, was
tumorigenesis in A/J mice (Johnson et al. 2008). about 4- to 12-fold more effective in reducing the
Kava treatment inhibited proliferation and cell viabilities of androgen receptor (AR)-
enhanced apoptosis in lung tumors, as shown by negative, HRPC cell lines DU145 and PC-3 than
a reduction in proliferating cell nuclear antigen AR-positive, hormone-sensitive prostate cancer
(PCNA), an increase in caspase-3, and cleavage cell lines LAPC4 and LNCaP, with minimal
of poly(ADP-ribose) polymerase (PARP). Kava effect on normal prostatic epithelial and stromal
treatment also inhibited the activation of nuclear cells. FKB induced apoptosis with an associated
factor kappaBNF-kappaB, a potential upstream increased expression of proapoptotic proteins:
mechanism of kava chemoprevention. In subse- death receptor-5, Bim and Puma and a decreased
quent studies, Johnson et al. (2011) reported that expression of inhibitors of apoptosis protein:
mice-fed diets containing kava at dosages of XIAP and survivin. Additionally, FKB syner-
1.25, 2.5, 5 and 10 mg/g of diet had 8.4, 6.6, gized with TNF-related apoptosis-inducing
4.3 ± 2.4 and 3.8 lung adenomas per mouse, ligand (TRAIL) for markedly enhanced induc-
respectively corresponding to a reduction of tion of apoptosis. Furthermore, FKB treatment of
31 %, 46 %, 65 % and 69 % in tumor multiplicity. mice bearing DU145 xenograft tumors resulted
They found that flavokawains A, B and C, from in tumor growth inhibition and increased Bim
kava, demonstrated greatly reduced chemopre- expression in tumor tissues. Flavokawain B sig-
ventive efficacies even at concentrations much nificantly inhibited the cell proliferation of
higher than their natural abundance, suggesting human adenoid cystic carcinoma in-vitro in a
that they alone were unlikely to be responsible dose-dependent manner that was associated with
for kava’s chemopreventive activity. Kava at all induced apoptosis and cell cycle G2-M arrest,
dosages and treatment regimens did not induce and the half maximal inhibitory concentration
detectable adverse effects, particularly with (IC50) of flavokawain-B treatment for 48 oursh
respect to liver. Specifically, kava treatment was estimated to be 4.69 μmol/L (Zhao et al.
showed no effect on liver integrity indicator 2011). Induction of apoptosis was by up-
enzymes or liver weight, indicating that kava regulation of Bim and down-regulation of Bcl-2
may be potentially safe for long-term chemopre- expression. Li et al. (2012) treated prostate can-
ventive application. Shaik et al. (2009) demon- cer (LNCaP, LAPC-4, 22Rv1, C4-2B, DU145
strated that kava suppressed nuclear factor-kappaB and PC-3) cell lines having different androgen
(NF-kappaB) activation in lung adenoma tissues, receptor (AR) expression and a transformed
potentially a mechanism responsible for kava’s prostate myofibroblast cell line (WPMY-1), with
chemopreventive activity. Methysticin was iden- a commercial kava extract, kavalactones (kawain,
tified as a potent NF-kappaB inhibitor in kava 5′6′-dehydrokawain, yangonin, methysticin) and
Piper methysticum 157

flavokawain B. The kava extract and flavokawain tration of flavokawain A (FKA), the predominant
B effectively down-regulated the expression of chalcone from kava, to UPII-SV40T transgenic
both the full-length AR and AR splice variants. mouse for 318 days inhibited the occurrence of
The kava extract and kavalactones accelerated high-grade papillary urothelial cell carcinoma, a
AR protein degradation, while flavokawain B precursor to invasive urothelial cancer, by 42.1 %
inhibited AR mRNA transcription via decreasing (Liu et al. 2013). A decreased expression of Ki67,
Sp1 expression and the binding of Sp1 to the AR survivin and X-linked inhibitor of apoptotic pro-
promoter. The kava root extract and flavokawain teins (XIAP) and increased expression of p27 and
B reduced tumor growth, AR expression in tumor DR5, and the number of terminal deoxynucleoti-
tissues and levels of serum PSA in the patient- dyl transferase-mediated dUTP nick end labeling
derived prostate cancer xenograft in mice. The (TUNEL)-positive apoptotic cells were observed
results suggested a potential usefulness of a safe in the urothelial tissue of FKA-fed mice. The
kava product or its active components for preven- results suggested a potential of FKA in prevent-
tion and treatment of advanced prostate cancer by ing the recurrence and progression of non-
targeting AR. muscle-invasive urothelial cell carcinoma.
Flavokawain B (FKB) potently inhibited the The results of animal studies by Triolet et al.
growth of synovial sarcoma cell lines SYO-I and (2012) suggested that kava may reduce colon
HS-SY-II through induction of apoptosis (Sakai cancer risk. After 14 weeks, rats fed the nonpolar
et al. 2012). Treatment with FKB increased cas- fraction of kava extract 12 days prior to, during
pase 8, 9 and 3/7 activity compared to vehicle- and after the administration of dimethylhydra-
treated controls, indicating that both extrinsic and zine, a colon-specific carcinogen, had a signifi-
intrinsic apoptotic pathways were activated. In cant reduction in precancerous lesions [aberrant
addition, FKB treatment of both cell lines resulted crypt (AC) foci (ACF)] as well as large (≥ 4 AC/
in increased mRNA and protein expression of ACF) sialomucin-only expressing foci, an indica-
death receptor-5 and the mitochondrial pro- tor of greater tumorigenic potential, compared to
apoptotic proteins Bim and Puma, while down- the control group (Triolet et al. 2012). Groups fed
regulating the expression of an inhibitor of the ethanolic kava extract and polar kava fraction
apoptosis, survivin in a dose-dependent manner. trended toward reductions in ACF and large
Treatment with flavokawain B (FKB), a novel sialomucin-only expressing foci. The combined
kava chalcone, preferentially inhibited the growth kava groups had significantly fewer total AC,
of uterine leiomyosarcoma (SK-LMS-1), endo- ACF, large ACF and large sialomucin-only
metrial adenocarcinoma (ECC-1) cells compared expressing foci compared to the control group.
to the non-malignant, human endometrium Histological examination found no hepatic
fibroblast-like (T-HESC) cell lines (Eskander lesions in animals consuming the kava diets.
et al. 2012). FKB treatment resulted in cell cycle Studies by Leitzman et al. (2014) demonstrated
arrest and a robust induction of apoptosis in the outstanding efficacy of kava in preventing
SK-LMS-1 and ECC-1 cell lines. 4-(methylnitrosamino)-1-(3-pyridyl)-1-butanone
In a recent study, flavokawain B, a kava chal- (NNK)-induced lung tumorigenesis in A/J mice
cone, inhibited the growth of human osteosar- with high selectivity for the initiation stage in
coma cell lines 143B and Saos-2 through G2/M association with the reduction of O6-methylguanine
cell cycle arrest and induction of apoptosis adduct in DNA. Kava treatments covering the ini-
involving both extrinsic and intrinsic pathways tiation stage reduced the multiplicity of lung
(Ji et al. 2013). Furthermore, migration and inva- adenomas by approximately 99 %. A minimum
sion ability was decreased by FKB in a dose- effective dose was not defined because kava at
dependent manner. The cytotoxicity profile two lower dosages (2.5 and 1.25 mg/g of diet)
showed FKB had significant lower side effects on were equally effective as 5 mg/g of diet in
bone marrow cells and small intestinal epithelial completely inhibiting lung adenoma formation.
cells compared with adriamycin. Oral adminis- Dihydromethysticin from kava completely
158 Piperaceae

suppressed tobacco carcinogen 4-(methyl- Aurora B. Altogether, the results elucidated the
nitrosamino)-1-(3-pyridyl)-1-butanone-induced anti-inflammatory mechanisms triggered by tra-
lung tumorigenesis and differentially reduced ditionally used chemopreventive kava com-
DNA damage in A/J mice (Narayanapillai et al. pounds for inflammatory diseases and asthma.
2014a). Flavokawain A (FKA) from kava induced Of the kava-derived compounds tested in-
apoptosis in two breast cancer cell lines, MCF-7 vivo, kavain was found to render mice immune to
and MDA-MB231, and inhibited the metastatic lethal doses of lipopolysaccharide (Pollastri et al.
process in-vitro (Abu et al. 2014). FKA also halted 2009). Kavain demonstrated promising pharma-
the migration and invasion process in ceutical properties, including good solubility and
MDA-MB231. high cell permeability, but pharmacokinetic
experiments in mice showed relatively rapid
clearance. Of the kavain analogues synthesized, a
Antiinflammatory Activity ring-opened analog of kavain inhibited TNF-
alpha secretion in THP-1 cell-based assay and
Five known kawapyrones, 5,6-dehydrokawain suppressed lipopolysaccharide-induced TNF-
(isolated yield: 0.12 %), dihydrokawain (0.35 %), alpha factor expression in the same cells, whereas
kawain (0.47 %), yangonin (0.22 %) and methys- the other compounds inhibited TNF-alpha secre-
ticin (5.66 %) along with a new kavapyrone, tion without affecting lipopolysaccharide-
7,8-epoxyyangonin (6) (0.0059 %) isolated from induced TNF-alpha factor levels, indicating a
the methanol extract of dried kava powder were potential divergence in mechanism of action.
tested for TNF-α (tumor necrosis factor-α) Flavokawain A, from kava extract, significantly
release assay from BALB/3 T3 cells treated with suppressed expression of iNOS and COX-2, as
okadaic acid, a tumor promoter (Hashimoto et al. well as the subsequent production of NO and
2003). 5,6-Dehydrokawain (desmethoxyyango- PGE2 in the LPS-stimulated RAW 264.7 cells
nin) (1) and yangonin (4) significantly inhibited (Kwon et al. 2013). Flavokawain A significantly
TNF-α release with IC50 values of 17 μM and inhibited LPS-induced activation of NF-κB and
40 μM; a potency as great as (−)-epigallocatechin AP-1 signaling pathways. The results suggested
gallate (EGCG) isolated from green tea extract. that flavokawain A may exert anti-inflammatory
Among the kavapyrones, dihydrokawain was responses by suppressing LPS-induced expres-
unique in showing the strongest inhibitory activ- sion of pro-inflammatory mediators via blockage
ity against TNF-α release in mice, but the weak- of NF-κB-AP-1-JNK/p38 MAPK signaling path-
est activity in the cells. Studies by Folmer et al. ways in the murine macrophages. Studies showed
(2006) found that 320 μM (flavokavain A), that a kavalactone derivative2′,6′-dichloro-5-
175 μM (flavokavain B) and 870 μM (kavain and methoxymethyl-5,6-dehydrokawain inhibited
dihydrokavain) inhibited both NF-kappa B-driven lipopolysaccharide-stimulated iNOS induction
reporter gene expression and TNF alpha-induced and NO production through activation of Nrf2
binding of NF-kappaB. Moreover, kavain and fla- signaling in BV2 microglial cells (Terazawa et al.
vokavains A and B treatment led to inhibition of 2013). The data suggested that the compound had
both inhibitors of kappaB (IkappaB) degradation a potential to reduce neuroinflammation as well
and subsequent translocation of p50 and p65 as oxidative stress in neurodegenerative diseases
NF-kappa B subunits from the cytoplasm to the through activation of Nrf2 signaling.
nucleus. Further, kinase selectivity screening
demonstrated that flavokavain A, but not kavain,
nor flavokavain B, inhibited the IkappaB kinase Central Nervous Sytem (CNS) Activity
(IKK) as well as PRAK (p38-regulated/activated
kinase), MAPKAP-K3 (MAPK-activated protein Animal studies showed that kava lactones altered
kinase 3), DYRK1A (dual-specificity tyrosine- neuronal excitation through direct interactions
phosporylated and regulated kinase 1A) and with voltage-dependent ion channels, giving rise
Piper methysticum 159

to kava’s muscle relaxant, anaesthetic, anxiolytic tration of kavain and dihydromethysticin in a

and anticonvulsive properties (Cairney et al. dose-dependent manner down to 22.2 and 33.6 %,
2002). Several isolated cases of psychotic and respectively. Intragastric administration of
severe dystonic reactions following kava use sug- (+)-dihydromethysticin in a single dose (100 mg/
gested that kava also had psychoactive proper- kg body weight) or chronic intragastric adminis-
ties, yet there was no conclusive evidence to-date tration of (±)-kawain (10.8 mg/kg body weight)
that kava interfered with normal cognitive pro- daily for 78 days to rats did not alter dopamine or
cesses. Although the psychotropic and neuropro- serotonin levels in the striatal or cortical brain
tective mechanisms of the rhizome are not well regions (Boonen et al. 1998).
elucidated, numerous studies had indicated that Dinh et al. (2001) found that the most potent
kava pyrones may exert their effects by activating binding inhibition was observed for kava leaf
several neurotransmitter systems, such as the extracts to GABAA receptors (GABA binding
adrenergic (Seitz et al. 1997b), mesolimbic dopa- site) with IC50 values of approximately 3 μg/ml,
minergic (Baum et al. 1998), gabaminergic whereas root extracts were less active with IC50
(Jussofie et al. 1994), glutamatergic (Schmitz values ranging from 5 μg/ml (Nene) to 87 μg/ml
et al. 1995; Schmidt and Ferger 2001; Gleitz (Mahakea). The leaf extracts generally contained
et al. 1996a,b) and serotonergic receptor systems lower amounts of the kavalactones than the root
(Walden et al. 1997b; Boonen et al. 1998). An extracts, indicating the existence of additional
extract of the rhizome containing 58 % kava substances responsible for these activities. Leaf
pyrones enhanced the binding of [3H]muscimol extracts also inhibited binding to dopamine D2,
to γ-aminobutyric acid-A receptors in a opioid (mu and delta) and histamine (H1 and H2)
concentration-dependent manner in rat hippo- receptors more potently than the corresponding
campus, amygdala and medulla oblongata in- root extracts with IC50 values ranging from 1 to
vitro (ED50 200–300 μmol/l) (Jussofie et al. 100 μg/ml vs. > or = 100 μg/l, respectively.
1994). Contrariwise the study by Davies et al. Significant differences in the potential of binding
(1992) found no significant interaction in-vitro or inhibition were also observed between cultivars.
in-vivo of a dichloromethane rhizome extract or Binding to serotonin (5-HT6 and 5-HT7) and
kava pyrones with γ-aminobutyric acid (A and B) benzodiazepine receptors was only weakly inhib-
or benzodiazepine receptor binding sites. Seitz ited by both root and leaf extracts of all four kava
et al. (1997b) showed that (+/−)-Kavain and cultivars. Among the kava lactones tested, only
(+)-kavain potently inhibited the uptake of yangonin exhibited affinity for the human recom-
[3H]-noradrenaline in synaptosomes prepared binant CB1 receptor ligand with a Ki = 0.72 μM
from the cerebral cortex and hippocampus of and selectivity vs. the CB2 receptor (Ki > 10 μM)
rats. Uptake of [3H]-noradrenaline was inhibited (Ligresti et al. 2012). None of the compounds
in the following order of potency: exhibited strong inhibitory effects on the two
(+/−)-kavain = (+)-kavain > (+)-methysticine, enzymes fatty acid amide hydrolase (FAAH) and
whereas none of the kava pyrones efficiently monoacylglycerol lipase (MAGL) analyzed. The
blocked the uptake of [3H]-serotonin. The results CB1 receptor affinity of yangonin suggested that
indicated a pyrone-specific non-stereo-selective the endocannabinoid system might contribute to
inhibition of the [3H]-noradrenaline uptake the complex human psychopharmacology of the
which might be responsible for or, at least, con- traditional kava drink and the anxiolytic prepara-
tribute to the psychotropic properties of kava tions obtained from the kava plant.
pyrones. Both kawain and dihydromethysticin
reduced the field potential changes induced by Anxiolytic Activity
the serotonin-1A agonist, ipsapirone, in the CA1
and CA3 areas of guinea-pig hippocampal slices In-Vitro Studies
in-vitro (Walden et al. 1997a. The ipsapirone Studies using hippocampal slice preparation of
response was reduced by extracellular adminis- guinea pigs by Walden et al. (1997a) suggested
160 Piperaceae

that single constituents of Piper methysticum kavain, dihydrokavain, methysticin, dihydrome-

may have additive actions; two kava components thysticin, yangonin or desmethoxyyangonin
kawain and dihydromethysticin may enhance the (30 mg/ml per kg for kava compounds) attenu-
effects of the anxiolytic serotonin-1A agonist ated separation-induced distress vocalizations
ipsapirone and activation of NMDA receptors and stress-induced analgesia in 8-day-old chicks
and/or voltage dependent calcium channels may (Smith et al. 2001). Dihydrokavain attenuated
be involved in the elementary mechanism of separation-induced distress vocalizations.
action of some kava-pyrones. A small dose of Studies showed that administration of a kava
kava extract (20 mg/kg body weight i.p.) caused preparation LI 150 to Wistar rats exhibited anxio-
changes in rat behaviour and concentrations of lytic like behaviour similar to diazepam (15 mg/
dopamine in the nucleus accumbens (Baum et al. kg p.o.) in the elevated plus maze test (Rex et al.
1998). Higher doses (120 mg/kg i.p.) increased 2002). Using the chick social separation-stress
the levels of dopamine. With respect to the indi- procedure as an anxiolytic bioassay, P. methysti-
vidual compounds, d,l-kawain induced in low cum samples containing 12.8–100.0 % total
doses a decrease in dopamine levels and in higher kavalactones in experiment 1 attenuated distress
amounts either an increase or no change in dopa- vocalizations in a concentration-dependent man-
mine concentrations. Yangonin resulted in a ner (Feltenstein et al. 2003). P. methysticum frac-
decrease of dopamine levels to below the detec- tions that contained the highest concentration of
tion limit and desmethoxyyangonin in an increase dihydrokavain attenuated distress vocalizations
of dopamine levels. Dihydrokawain, methysticin in a manner equivalent to that of chlordiazepox-
and dihydromethysticin did not produce any sig- ide in separate experiments. Kava extract samples
nificant changes of dopamine levels. d,l-kawain and fractions that possessed anxiolytic properties
caused a decrease in 5-HT concentrations. Some did not possess the sedative properties found in
of the other kavapyrones affected 5-HT levels as chlordiazepoxide. Collectively, the findings sug-
well. The results suggested that the relaxing and gested that dihydrokavain may be necessary and
slightly euphoric actions may be caused by the sufficient in mediating the anxiolytic properties
activation of the mesolimbic dopaminergic neu- of P. methysticum extract.
rones. Changes in the activity of 5-HT neurones Kava extract produced statistically significant
could explain the sleep-inducing action. Studies dose-dependent anxiolytic-like behavioral
by Langosch et al. (1998) found (+/−)-kavain to changes in BALB/cByJ inbred mice in both the
be an effective drug in modulating excitatory sig- mirrored chamber avoidance and elevated plus-
nals in the hippocampus of guinea pigs. In their maze assays of anxiolysis (Garrett et al. 2003).
paper, Nowakowska et al. (1998) concluded that ED50 values for kava-induced increases in time
kava-kava may be a useful alternative for syn- spent inside the mirrored chamber and on the
thetic anxiolytics. open arms of the plus maze were 125 mg/kg and
88 mg/kg, respectively. Kava extract also caused
Animal Studies a profound decrease in locomotor activity
Aqueous kava root extracts were shown to have a (ED(50) of 172 mg/kg). Flumazenil, a competi-
depressant action on the central nervous system, tive benzodiazepine receptor antagonist, blocked
evidenced by depression of spontaneous motor both the anxiolytic and sedative effects of diaze-
activity, marked reduction in irritability of rats pam, but had no effect on kava’s behavioral
having bilateral septal lesions, inhibition of the actions. The results showed that kava anxiolytic
conditioned avoidance response in rats and and sedative effects were not mediated through
blockade of EEG arousal patterns in cats the benzodiazepine binding site on the GABA(A)
(Furgiuele et al. 1965). Both chlordiazepoxide receptor complex. In animal studies, Bruner and
and kava extract (containing 30 % kavalactones), Anderson (2009) found dose-dependent substitu-
Piper methysticum 161

tion of the anxiolytic chlordiazepoxide (CDP) by nificantly reduced anxiety symptoms after
kava extracts. Partial substitution of kava extract 1 week; no adverse effects were noted during the
for CDP suggested that kava may share a mecha- 4-week treatment (Kinzler et al. 1991). In a one
nism of action similar to CDP, but was less 6-month, double-blind study compared kava was
potent. found to be just as effective as two standard anxi-
ety benzodiazepine drugs (oxazepam and bro-
Clinical Studies mazepam) in 174 people with anxiety symptoms
Two placebo-controlled trials investigated the (Woelk et al. 1993). Improvement in Hamilton
effect of standardized Kava extracts in women Anxiety Scale (HAMA) scores was about the
with climacteric psychosomatic disturbances same in all groups. A 4 weeks, randomized,
(Warnecke et al. 1990; Warnecke 1991). In the placebo-controlled double-blind study of 58
first study involving 40 such women, kava extract patients also assessed the efficacy of WS 1490
(200–400 mg extract = 30–60 mg kava pyrones) extract containing 70 % kava pyrones for the
for 8 weeks was found to be superior to the pla- treatment of anxiety of non-psychotic origin
cebo using the Kuppermann Index and Anxiety (Lehmann et al. 1996). The Hamilton Anxiety
Status Index. In the second randomized, placebo- Scale (HAMA) overall score of anxiety symp-
controlled double-blind study, two groups each tomatology revealed a significant reduction in the
containing 20 patients with climacteric-related kava group compared with the placebo group
symptomatology were treated for a period of 8 after 1 week of treatment. The results of the sec-
weeks with kava WS 1490 extract 3 × 100 mg/day ondary target variables were in agreement with
or a placebo preparation. The Hamilton Anxiety the HAMA score and demonstrated the efficacy
Scale (HAMA) overall score of anxiety symp- of WS 1490 in patients with anxiety disorders. In
tomatology revealed a significant difference in another randomized placebo-controlled 25-week
the kava-receiving group compared to the pla- outpatient trial of 101 outpatients suffering from
cebo group already after only 1 week of treat- anxiety of non-psychotic origin (Diagnostic and
ment. The course of such further parameters as Statistical Manual of Mental Disorders (DSM)
depressive mood (DSI), subjective well-being -III-R criteria: agoraphobia, specific phobia, gen-
(patient diary), severity of the disease (CGI) and eralized anxiety disorder and adjustment disorder
the climacteric symptomatology (Kuppermann with anxiety), treatment with kava-kava extract
Index and Schneider scale) over the overall WS 1490 was found to be superior over placebo
period of treatment demonstrated a high level of from week 8 onwards in the Hamilton Anxiety
efficacy of kava extract WS 1490 in neurovegeta- Scale (HAMA) (Volz and Kieser 1997). WS 1490
tive and psychosomatic dysfunctions in the cli- was also found to be superior with respect to the
macteric, associated with very good tolerance of secondary outcome variables. The results sup-
the preparation. A randomized, double-blind ported WS 1490 as a treatment alternative to tri-
pilot study investigated the effects of the extract cyclic antidepressants and benzodiazepines in
containing 15 % kava pyrones in 59 patients with anxiety disorders. In an outpatient observation
pre-operative anxiety (Bhate et al. 1989). study of 52 outpatients suffering from anxiety of
Although improvements in mood were observed nonpsychotic origin, the treatment of kava extract
using a psychostatus score, only two doses of the was found to be as an effective and safe alterna-
extract (equivalent to 60 mg kava pyrones daily) tive to antidepressants and tranquilizers in anxi-
were administered, and thus the clinical signifi- ety disorder without the tolerance problems
cance of this study was questionable. associated with benzodiazepines (Scherer 1998).
In a placebo-controlled double-blind study of The results of a randomised study of 40 women
29 patients with anxiety syndromes, treatment in physiological or surgical menopause for the
with 100 mg of WS 1490 three times a day sig- past 1–12 years showed that the association of
162 Piperaceae

hormone replacement therapy (HRT) and kava- the same extract. WS 1490 was well tolerated,
kava extract may represent an excellent therapeu- with no influence on liver function tests and only
tic tool for the treatment of women in stabilized one trivial adverse event (tiredness) attributable
menopause, in particular those suffering from to the study drug. In a 3-week placebo-controlled,
anxiety and depression, given that kava-kava double-blind crossover trial that recruited 60
therapy accelerated the resolution of psychologi- adult participants with 1 month or more of ele-
cal symptoms without diminishing the therapeu- vated generalized anxiety, administration of 5
tic action of estrogens on organic disease, such as kava tablets per day containing 250 mg of kava-
osteoporosis and cardiovascular disease (De Leo lactones/day, produced significant anxiolytic and
et al. 2000). antidepressant activity and raised no safety con-
Malsch and Kieser (2001) conducted a 5-week cerns at the dose and duration studied (Sarris
randomised, placebo-controlled, double-blind et al. 2009b). Kava appeared equally effective in
study to investigate the efficacy of kava-kava spe- cases where anxiety was accompanied by depres-
cial extract WS1490 in 40 patients with non- sion. In another 3-week placebo-controlled,
psychotic nervous anxiety, tension and double-blind, cross-over trial involving 60 adult
restlessness states. The study confirmed the anx- participants (18–65) with elevated stable anxiety
iolytic efficacy and good tolerance of WS1490 and varying levels of depressive symptoms, kava
over placebo and showed that a further symptom preparation exerted significant anxiolytic effects
reduction was possible after a change over from and appeared safe (Sarris et al. 2009a). In a ran-
benzodiazepine treatment. Studies by Watkins domized, placebo-controlled, double-blind study
et al. (2001) suggested that kava might exert a of 22 moderately anxious adults aged between 18
favourable effect on reflex vagal control of heart and 65 years, kava administration was found to
rate in generalized anxiety disorder patients. have no negative effect on cognition, whereas a
Significantly more patients treated with kava reduction in alertness occurred in the oxazepam
showed improved baroreflex control of heart rate condition (Sarris et al. 2012). Genetic analyses
(BRC) compared to the placebo group. Further, provided tentative evidence that noradrenaline
the magnitude of improvement in BRC was sig- (SLC6A2) transporter polymorphisms may have
nificantly correlated with the degree of clinical an effect on response to kava. They concluded
improvement. In a 3-month randomized prospec- that acute “medicinal level” doses of the particu-
tive open study of perimenopausal women, the lar kava cultivar in naive users did not provide
administration of kava-kava plus calcium elicited anxiolytic activity, although the phytomedicine
an improvement of mood, particularly of anxiety also appeared to have no negative effects on
(Cagnacci et al. 2003). Gastpar and Klimm cognition.
(2003) investigated the efficacy and tolerability In a more recent 6-week double-blind, ran-
of 150 mg/day kava special extract WS 1490 ver- domized, placebo-controlled study of 75 partici-
sus placebo in a randomized, placebo-controlled, pants with generalized anxiety disorder (GAD)
double-blind multicenter study in 41 adult, male and no comorbid mood disorder, treatment with
and female out-patients suffering from neurotic kava produced a significant reduction in anxiety
anxiety wherein patients received 3 × 1 capsule of compared with the placebo group with a moder-
50 mg/day WS 1490 or placebo for 4 weeks, fol- ate effect size (Sarris et al. 2013b). Among par-
lowed by 2 weeks of observation without study- ticipants with moderate to severe Diagnostic and
specific treatment. The results showed consistent Statistical Manual of Mental Disorders-diagnosed
advantages for WS 1490 over placebo in several GAD, this effect was larger. The data suggested
psychiatric scales and indicated significant that standardized kava may be a moderately
improvements in the patients’ general well-being, effective short-term option for the treatment of
the differences versus placebo were not as large GAD. Furthermore, specific γ-aminobutyric acid
as in previous trials which employed 300 mg/d of (GABA) transporter polymorphisms appeared to
Piper methysticum 163

potentially modify anxiolytic response to kava. 1959; Meyer 1962; Meyer and May 1964). Frater
In another 6-week, double-blind, randomized (1958) demonstrated that a thin paste of kava
controlled trial (n = 75) involving chronic admin- powder applied to the lip mucous membrane pro-
istration of kava or placebo for participants with duced a slightly burning sensation and a feeling
generalized anxiety disorder, no significant dif- of numbness when some kava roots were chewed
ferences were found across groups for liver func- for 15 min found the degree of anaesthesia was
tion tests nor were there any significant adverse greater. Steinmetz (1960) described the narcotic
reactions that could be attributed to kava (Sarris effects of kava as a paralysis of the nervous sys-
et al. 2013c). Interesting, kava significantly tem, through reduction of spinal rather than cere-
increased female’s sexual drive compared to pla- bral activity, followed by muscular stimulation
cebo on a sub-domain of the Arizona Sexual and then paralysis particularly affecting the lower
Experience Scale (ASEX), with no negative limbs. Also noted was a reduction of the cardiac
effects seen in males. Further, it was found that rhythm followed by stimulation and slowing
there was a highly significant correlation between down of respiration. Steinmetz also stated that
ASEX reduction (improved sexual function and ingestion of a strong dose would affect eye vision,
performance) and anxiety reduction in the whole causing pupil dilation and inducing photophobia.
sample. A gel containing 6 % kava root crude extract
called kavacaine was found to have potent topical
Meta-analysis/Review Studies anaesthetic effect (Morse and Sharma 2005).
In a meta-analysis of 11 randomised, controlled Administration of dihydromethysticin to mice
trials (RCTs), i.e. trials with a randomised gen- potentiated pentobarbital-induced sleeping time
eration of allocation sequences, and conducted by 400 %, while dihydrokawain, yangonin and
placebo-controlled and double-blind, i.e. trials kawain were only moderately active (150–235 %)
with blinding of patients and care providers and (Klohs et al. 1959). Both dihydrokawain and
involving a total of 645 participants, Pittler and dihydromethysticin exhibited analgesic effects
Ernst (2003) found that kava extract appeared to when administered intraperitoneally to rats
be an effective symptomatic treatment option for (140 mg/kg body weight), as determined by an
anxiety compared to placebo. The data available increase in tail-flick reaction times (Brüggemann
from the reviewed studies suggested kava to be and Meyer 1962). Meyer (1962) found that
relatively safe for short-term treatment (1–24 administration of 150 mg/kg of hexobarbital
weeks), although more information was required. sodium to white mice afforded 2 h of sleep.
In a comprehensive review of efficacy, safety and However, pretreament with 240 mg/kg dihydro-
psychopharmacology, Sarris et al. (2011) found methysticin induced 72 h of sleep. Meyer and
that current evidence supported the use of kava in May (1964) found all the kava lactones acted as
treatment of anxiety with a significant result local analgesics. Most of the kava lactones inhib-
occurring in four out of six studies reviewed. It ited frog heart contraction. These actions were
was suggested that more studies were required to comparable to cocaine which also protected
assess comparative efficacy and safety (on the against ventricular fibrillation through its local
liver, cognition, driving and sexual effects) ver- anaesthetic effect. Dihydroymethysticin and
sus established pharmaceutical comparators. dihydrokavain had been shown to intensify sleep-
inducing effects (Hänsel 1968). Both compounds
were found to be comparable to the drug dimeth-
Sedative/Analgesic/Anaesthetic/ ylaminophenazone in producing an analgesic
Anti-insomnia Activities effect in the drinker. The analgesic effect was
found to be stronger than aspirin but weaker then
The local anaesthetic activity of kava was first morphine. Kavain had been shown to produce
observed by Lewin (1886). Kava had been local aesthetic effect that was comparable to
reported to possess and to potentiate and prolong cocaine and do not produce any toxicity in the
barbiturate narcosis (Frater 1958; Klohs et al. tissues (Kretzschmar and Meyer 1969).
164 Piperaceae

The extent of frog sartorius muscle paralysis been reported to be taken in conjunction with
induced by kava was similar in both directly and alcoholic drinks. Psychopharmacological effects
indirectly stimulated mouse hemidiaphragms of kava had also been reported and the mood-
(Singh 1983). Intracellular recordings from frog altering effects of kavalactones had been
sartorius muscles showed that kava depressed the described as hypnotic (Schultes and Hoffman
amplitude of both miniature end-plate potentials 1992). The pharmacological properties of kava-
(mepps) and end-plate potentials (epps) but had lactones were comparable to those of benzodiaz-
no effect on the frequency of mepps. Kava greatly epines, however, kavalactones bind very weakly
prolonged the duration of mepps and epps and to the gamma-aminobutyric acid (GABA) and
also slowed and depressed directly elicited mus- benzodiazepine (Bilia et al. 2002). It was pro-
cle action potentials. The neuromuscular block- posed that N-methyl-d-aspartate receptors and/or
ade produced was poorly reversed by calcium voltage dependent calcium channels may also be
and by neostigmine. It was concluded that kava involved in the elementary mechanism of action.
caused paralysis by mechanisms similar to local Only weak activity of kava resin was observed on
anaesthetics. Both the intoxicating beverage kava GABAA binding sites washed synaptosomal
and the lipid soluble (kava resin) extracts showed membranes prepared from rat brain and this was
analgesic effects in both tail immersion and abolished by extraction of the membranes with
abdominal constriction in mice (Jamieson and Triton X-100, suggesting that lipid soluble com-
Duffield 1990b). Eight purified pyrones from the ponents were involved (Davies et al. 1992). No
lipid soluble extract were also tested for activity effects were observed on GABAB binding sites
in the tail immersion test, and kawain, dihydro- in rat brain membranes in-vitro. Kava resin and
kawain, methysticin and dihydromethysticin pyrones exerted some weak effects on benzodiaz-
were found to be very effective in producing epine binding in-vitro but this did not correlate
analgesia. Naloxone, in doses which inhibited with pharmacological activity. In addition, in ex-
morphine-induced analgesia in both tests, was vivo studies, no effects were observed on [3H]
indicating that analgesia produced by kava diazepam binding to brain membranes prepared
occurred via non-opiate pathways. The synthe- from mice in which selected kava constituents
sized analogues of synthetic kavain, were injected intraperitoneally, whereas similarly
6-(4-fluorophenyl)-4-methoxy-5,6-dihydropyran- administered diazepam (5 mg/kg) inhibited [3H]
2-one exerted potent and dose-dependent diazepam binding by greater than 95 %. Similar
analgesic activity, inhibiting abdominal constric- lack of activity was observed in in-vivo binding
tions caused by acetic acid in mice, and being studies; injection of kava resin failed to influence
more active than some reference drugs (Kormann the CNS binding of the benzodiazepine-receptor
et al. 2012). It also presented activity in the other ligand [3H]Ro15-1788 injected into mice prior to
models of pain, with the exception of the hot sacrifice. The pharmacological activities of kava
plate test and the measurement of motor resin and pyrones did not appear to be elucidated
performance. by any significant interaction with GABA or ben-
The lipid soluble extract of the psychoactive zodiazepine binding sites. Studies by Jussofie
beverage kava was found to have hypnosedative et al. (1994) indicated that one way kavapyrone-
properties which could be measured by the length enriched kava extract might mediate sedative
of time that the righting reflex was lost (Jamieson effects in-vivo was through effects on gamma-
and Duffield 1990a). Ethanol and the lipid solu- aminobutyric acid (GABA) receptor binding.
ble extract (kava resin) was shown greatly to The sedative effect was demonstrated using
increase each others’ hypnotic action in mice. membrane fractions obtained from target brain
Ethanol also increased the toxicity of kava mark- centers of kavapyrone action: hippocampus
edly. This interaction of kava and alcohol pos- (HIP), amygdala (AMY) and medulla oblongata
sessed important clinical and social consequences (MED), and from brain centers outside the main
since, in contrast to traditional usage, kava had kavapyrone effects as frontal cortex (FC) and cer-
Piper methysticum 165

ebellum (CER). At a kavapyrone concentration of do not produce any toxicity in the tissues
500 μM, the order of enhancement in binding (Kretzschmar and Meyer 1969). Examination of
sites (Bmax) was HIP = AMY > MED > FC > CE EEG recordings during sleep of healthy volun-
R. When kavapyrones were included together teers given a single dose of 300 mg extract
with pentobarbital or 3-alpha-hydroxy-5-alpha-- (equivalent to 210 mg kava pyrones) showed an
pregnane-20-one (HPO), the two classes of com- increased sleep spindle density of 20 % and an
pounds produced a synergetic effect on [3H] increase in slow-wave sleep (i.e. deep sleep), but
muscimol binding. the rapid eye movement phase was not sup-
In a double-blind, placebo-controlled study, pressed (Emser and Bartylla 1991). Daily doses
the encephalotropic and psychotropic effects of of 300 or 600 mg extract (equivalent to 210 or
kavain-a synthetic kava plant derivative as com- 420 mg kava pyrones), respectively, for 1 week
pared with clobazam were investigated in 15 nor- increased the beta/alpha index typical for the
mal volunteers, utilizing EEG brain mapping, pharmaco-EEG profile of anxiolytics without the
psychometric and psychopysiological analyses sedative-hypnotic effects associated with benzo-
(Saletu et al. 1989). Brain maps of drug induced diazepines. The increase in beta activity was
pharmaco-EEG changes (pharmaco-EEG maps) most marked in the beta2 range (Johnson et al.
demonstrated that kavain exerted a significant 1991). In two studies, administration of 300 mg
action on the human brain function as compared kava extract WS 1490 (equivalent to 210 mg kava
with placebo characterized by a dose-dependent pyrones) daily for either 8 or 14 days to healthy
increase of delta, theta and alpha 1 activity while volunteers, taken with or without ethanol, had no
alpha 2, beta activity and the centroid of the total potentiating or additive effect and had no influ-
activity decreased. These findings were indica- ence on the safety-related performance of healthy
tive of a sedative effect which was, however, in volunteers (Herberg 1991, 1993). In another
type quite different from that of the study administration of kava extract standardized
1,5-benzodiazepine. Further, 200 mg kavain to contain 30 % kava pyrones, bromazepam or a
induced with a decrease of delta and beta activity combination of extract and bromazepam, safety-
and an increase of alpha activity and of total related performance remained unaffected in
power also vigilance promoting effects. Kavain healthy volunteers treated daily with 400 mg
improved the noopsyche as compared with pla- extract (equivalent to 120 mg kava pyrones for 14
cebo in all three doses as there was a significant days), whereas it was impaired after treatment
improvement in intellectual performance (Pauli with bromazepam (9 mg daily) or the extract/bro-
test), attention, concentration, reaction time and mazepam combination (Herberg 1996). No dif-
motor speed (rigidity test), while opposite find- ferences were observed following treatment with
ings were observed after 30 mg clobazam. In bromazepam or the combination, indicating that
regard to thymopsychic variables such as drive, the extract did not have an additive effect when
wakefulness, affectivity, mood and well-being, given in combination with bromazepam. In a
200 mg kavain produced an improvement as separate, randomized, double-blind crossover
compared with placebo while 600 mg kavain pro- study involving 12 healthy volunteers, adminis-
duced sedation as did 30 mg clobazam. tration of daily single doses of a kava extract
Dihydroymethysticin and dihydrokavain had standardized to contain 30 % kava pyrones
been shown to intensify sleep-inducing effects (400 mg extract containing 120 mg kava pyrones)
(Hänsel 1968). Both compounds were found to was compared with daily single doses of diaze-
be comparable to the drug dimethylaminophena- pam (10 mg) or a placebo in a 7-day trial (Gessner
zone in producing an analgesic effect in the and Cnota 1994). Changes in EEG recordings
drinker. The analgesic effect was found to be and psychometric test results showed no evidence
stronger than aspirin but weaker than morphine. of a decrease in vigilance in the group treated
Kavain had been shown to produce local aes- with the extract. Following administration of
thetic effect that was comparable to cocaine and kava-kava extract at a dose of 300 mg/kg, a sig-
166 Piperaceae

nificant shortening of the sleep latency in sleep- the lipid-soluble extract (kava resin) reduced
disturbed rats was observed while no effects were amphetamine-induced hypermotility (Duffield
observed on the total waking and non-rapid eye et al. 1989b). Aqueous kava extract in i.p. doses
movement (non-REM) sleep time (Shinomiya of 30 mg/kg to 500 mg/kg had no effect on con-
et al. 2005). In contrast, flunitrazepam showed a ditioned avoidance responses. At or below
significant shortening in sleep latency, decrease 100 mg/kg i.p., kava resin also failed to modify
in total waking time and increase in total non- the number of conditioned avoidance responses
REM sleep time. Although the effects of fluni- obtained. However, 125 mg/kg of resin signifi-
trazepam were antagonized by the benzodiazepine cantly reduced the number of conditioned avoid-
receptor antagonist flumazenil, the effect of kava- ance responses by 18 %. Increasing the dose of
kava extract was not antagonized by flumazenil. kava to 150 mg/kg caused ataxia and sedation. A
Kava-kava extract showed a significant increase minimally effective daily dose (50 mg/kg) of the
in delta activity during non-REM sleep in sleep- aqueous kava extract for 3 days was sufficient to
disturbed rats, whereas a significant decrease in produce tolerance to a test dose of 150 mg/kg,
delta power during non-REM sleep was observed which was close to the ED50 (Duffield and
with flunitrazepam. The results showed that Jamieson 1991). Kava resin decreased spontane-
Kava-kava had not only hypnotic effects but also ous motility and caused a loss of muscle control.
sleep quality-enhancement effects. A minimally effective daily dose of kava resin
Two fractions, F1 and F2, from kava extract (100 mg/kg) did not produce tolerance to the
were found to decrease spontaneous motor activ- above effects of a weekly test dose of kava resin
ity in doses which did not alter forced motor (166 mg/kg) within 7 weeks. Increasing the dose
activity of mice (O’Hara et al. 1965). Fraction to 150 mg/kg twice daily caused partial tolerance
F2, dihydromethysticin, desmethoxy-yangonin to occur within 3 weeks, but very little further
and kawain exhibited potent antiserotonin activ- tolerance developed over the ensuing 2-week
ity on the isolated rat uterus, whereas F1 appeared period. Intraperitoneal administration of an
to be devoid of antiserotonin activity. F1, F2 and extract of kava rhizome (equivalent to 50–100 mg
dihydromethysticin did not alter serotonin brain kava pyrones/kg body weight) or (±)-kawain, a
levels in mice. Intraperitoneal (i.p.) adminstra- synthetic kava lactone (10–50 mg/kg body
tion of an aqueous extract of kava (62.5 mg/kg weight), reduced muscle tone in cats (Holm et al.
body weight) caused a loss of spontaneous activ- 1991). Intraperitoneal administration of kava rhi-
ity without loss of muscle tone in mice (Jamieson zome (equivalent to 50–100 mg kava pyrones/kg
et al. 1989). No hypnotic effect was seen, but body weight) to cats had a significant effect on
some analgesia was produced. The anticonvul- EEG recordings, inducing high-amplitude delta
sant effect against strychnine was very slight and waves, spindlelike formation and continuous
there was no evidence of local anesthetic action. alpha- or beta-synchronization in amygdala
There was a slight anti-apomorphine effect and recordings. Hippocampal responses, following
tetrabenazine-induced ptosis was decreased. The stimulation of the amygdala nucleus, increased
lipid-soluble extract (kava resin) also decreased significantly in amplitude in cats treated intra-
spontaneous motility and markedly reduced peritoneally with the rhizome extract or
motor control (Jamieson et al. 1989). Hypnosis, (±)-kawain.
determined by loss of righting reflex, was pro-
duced, analgesia was marked, and a local anes-
thetic action was evident. Kava resin also Anticonvulsant Activity
decreased apomorphine-induced hyper-reactivity
and partially reversed tetrabenazine-induced pto- Intraperitoneal administration of an aqueous
sis in rats. Kava resin produced a greater range of extract (300 mg/kg body weight) or a chloroform
pharmacological actions than the aqueous extract, extract (140 mg/kg body weight) of the rhizome
and the latter was orally inactive in mice and rats. to mice-inhibited strychnine-induced convul-
The aqueous, pyrone-free extract from kava and sions (Klohs et al. 1959). The anticonvulsant
Piper methysticum 167

activity of methysticin and other kava pyrones slices containing the hippocampus and the ento-
against electroshock- and chemically induced rhinal cortex, methysticin in a concentration
seizures has been demonstrated in mice and rats range of 10–100 μM reversibly blocked all types
(Keller and Klohs 1963; Meyer and Meyer-Burg of epileptiform activity (Schmitz et al. 1995).
1964; Kretzschmar and Meyer 1969; Kretzschmar While responses to alvear stimulation were
et al. 1970). Intraperitoneal administration of largely unaffected by methysticin, the responses
dihydromethysticin and dihydrokawain inhibited to a paired pulse stimulus to stratum radiatum
electroshock-induced seizures at doses of 25 and were depressed over the whole range of tested
60 mg/kg body weight, respectively, in mice stimulus intervals. The findings suggested that
and rats (Meyer and Meyer-Burg 1964). methysticin had effects on different patterns of
Dihydrokavain and dihydromethysticin inhibited epileptiform activity possibly by interfering with
muscular contractions and the effect was found to processes responsible for frequency potentiation.
be comparable to those of synthetic products Both (±)-kawain and methysticin inhibited
such as phenobarbital (Kretzschmar and Meyer voltage-dependent sodium channels in rat CA1
1969). Dihydromethysticin was reported to have hippocampal neurons in-vitro (1–400 μmol/l)
strong anticonvulsant action with the capability (Magura et al. 1997). In cultured dorsal root gan-
of inhibiting convulsions caused by strychnine in glion cells derived from neonatal rats,
animals (Kretzschmar et al. 1970). Both Schirrmacher et al. (1999) found that (+/−)-kavain
compounds were reportedly more superior to reduced currents through voltage-activated Na+
benzodiazepine as muscle relaxants. It had and Ca2+ channels.
been suggested that the anti-epileptic action of Kava pyrones were reported to exert effects on
dihydryomethysticin may be used to treat schizo- neuronal transmission and transmembraneous
phrenia. Isolated kavain, methysticin, dihydro- cation currents similar to established mood stabi-
methysticin and yangonin showed strong lizers like carbamazepine, valproate and
centrally mediated muscle relaxing activity in lamotrigine (Grunze et al. 2001). Findings of
rabbits and yangonin was the most potent kava- studies suggested that (i) kava pyrones had a
lactone (Kretzschmar et al. 1971). Studies had weak Na+ antagonistic effect that may contribute
shown kava pyrones to be anticonvulsive (Gleitz to their antiepileptic properties; (ii) they had pro-
et al. 1996b). The anticonvulsive action of nounced l-type Ca2+ channel antagonistic proper-
(+/−)-kavain was estimated from its properties on ties and acted as a positive modulator of the early
stimulated synaptosomes and Na+ channel recep- K+ outward current; two actions of importance
tor sites. The results suggested an interaction of for mood stabilization; (iii) kava pyrones had
(+/−)-kavain with voltage-dependent Na+ and additive effects with the serotonin-1A agonist
Ca2+ channels, thereby suppressing the ipsapirone probably contributing to their anxio-
4-aminopyridine-induced increase in [Na+]i, lytic and sleep-inducing effects; and (iv) kava
[Ca2+]i and the release of endogenous glutamate. pyrones showed a distinct pattern of action on
(+/−)-Kavain specifically and rapidly inhibited glutamatergic and GABAergic transmission
veratridine-activated voltage-dependent Na+- without affecting long-term potentiation. It was
channels in synaptosomes prepared from rat found that kava pyrones exhibited a profile of cel-
cerebral cortex (Gleitz et al. 1995). Further lular actions that showed a large overlap with
they found that (+/−)-kavain inhibited the several mood stabilizers, especially lamotrigine.
veratridine-induced and KCl-induced increase in
intracellular Ca2+ and glutamate-release of rat
cerebrocortical synaptosomes (Gleitz et al. Neuroprotective Activity
1996a). The results suggested that (+/−)-kavain
at concentrations sufficient to block Na+ channels The neuroprotective effects of an acetone extract
completely, moderately inhibited the non- of kava rhizome and kava pyrones had been dem-
inactivating Ca2+ channels located on mammalian onstrated both in-vivo and in-vitro. Administration
presynaptic nerve endings. In rat temporal cortex of kava extract (orally) and its constituents
168 Piperaceae

methysticin and dihydromethysticin (intraperito- effects of oxazepam and kava root extract on
neally) protected against brain tissue against behavior and event-related potentials (ERPs) in
ischemic damage in mice and rats (Backhauss a recognition memory task in twelve healthy
and Krieglstein 1992). The kava extract (150 mg/ volunteers. Oxazepam led to a reduction of a
kg, 1 h before ischemia) diminished the infarct negative component in the 250–500 ms range
area in mouse brains and the infarct volume in rat for both old and new words and to a reduction of
brains. Methysticin, dihydromethysticin (both 10 the old/new difference in the ERP associated
and 30 mg/kg, 15 min before ischemia) and the with a significantly worse recognition rate.
reference substance memantine (20 mg/kg, Kava, on the other hand, showed a slightly
30 min before ischemia) significantly reduced the increased recognition rate and a larger ERP dif-
infarct area in mouse brains. The standardized ference between old and new words Heinze
extract also protected against neuronal damage in et al. (1994) investigated the effects of oxaze-
cultured neurons from chick embryo cerebral pam and a standardized extract of kava roots
hemispheres (Backhauss and Krieglstein 1992). (WS1490) on reaction time and event-related
Kava pyrones exhibited neuroprotective and potentials (ERPs) in a visual search paradigm
“recovery-supporting” effects on neurological using a double-blind design in young, healthy
deficits after cerebral infarction in rats (Kleiser males. Significant effects of oxazepam were
et al. 1998). obtained in a number of psychometric tests.
Administration of the lower dosages of Oxazepam led to a reduction of the amplitude of
(+/−)-kavain (50 and 100 mg/kg) to male the parietal N1, frontal N2, posterior contralat-
C57BL/6 mice afforded only a nonsignificant eral N2 and occipital P3 components. WS 1490
attenuation of MPTP (1-methyl-4-phenyl- was associated with a greater posterior N1, pos-
1,2,3,6-tetrahydropyridine)-induced dopamine terior contralateral N2 and occipital P3. In a
depletion, but a high dosage of (+/−)-kavain double-blind randomized placebo-controlled
(200 mg/kg) significantly antagonized the dopa- trial involving healthy volunteers, Thompson
mine depletion to 58.93 % of saline control val- et al. (2004) found the intake of a single dose of
ues (Schmidt and Ferger 2001). The kava extract (300 mg; p.o.) led to an increase in
MPTP-induced decrease of tyrosine hydroxylase- state cheerfulness, while the herbal medicine
immunoreactivity as well as the loss of nigral did not influence state seriousness and bad
neurons was completely prevented by mood. The mood-elevating effects of kava were
(+/−)-kavain (200 mg/kg). They concluded that most prominent in trait cheerful subjects, indi-
MPTP metabolism was not influenced by cating that trait cheerfulness moderated the
(+/−)-kavain and postulated the antiglutamater- drug-induced increase in cheerful mood.
gic effects of (+/−)-kavain for its protective Further, Kava improved the accuracy and the
effects against MPTP toxicity. They asserted that speed of performing the partial report and the
kavain may be a novel candidate for further pre- item recognition task, indicative of a beneficial
clinical studies in animal models of Parkinson’s effect of kava on visual attention and short-term
disease and other disorders with glutamatergic memory retrieval, respectively.
overactivity. In a systematic review of clinical trials on the
effects of kava on cognition, La Porte et al. (2011)
found the majority of evidence from 10 human
Cognitive Activity clinical trials suggested that kava had no repli-
cated significant negative effects on cognition.
Kava was found to have no effect on the reaction One acute study found that kava significantly
times or errors of two groups of undergraduates improved visual attention and working memory
who consumed doses thought usual (Experiment processes while another found that kava increased
1) and doses much greater (Experiment 2) than body sway. One chronic study found that kava
those associated with social functions (Russell significantly impaired visual attention during
et al. 1987). Münte et al. (1993) investigated the high-cognitive demand.
Piper methysticum 169

In a randomized, placebo-controlled, double- the K+ channel by barium chloride (1 mM) or

blind study of 22 adults aged between 18 and 65 4-aminopyridine (0.3 mM). After pre-incubation
years, random administration of an acute medici- with 1 μM nifedipine, carbachol (1 μM) evoked
nal dose of kava (180 mg of kavalactones), oxaz- 18.2 % of contraction at control (i.e. prior pre-
epam (30 mg) or placebo 1 week apart in a incubation with nifedipine). This remaining
crossover design showed that the medicinal dose response was completely abolished by high con-
of kava containing 180 mg of kavalactones did centrations of (±)-kavain (400 μM). After treat-
not impair driving ability, whereas 30 mg of ment of the longitudinal ileum strips with
oxazepam showed some impairment (Sarris et al. pertussis toxin, carbachol (1 μM) evoked 27.0 %
2013a). of the control response in untreated ileum. These
contractions were also blocked by (±)-kavain
(400 μM). However, (±)-kavain had no effect on
Antispasmodic Activity the caffeine-induced (20 mM) contractions of
ileum strips. Moreover, it failed to affect Ca2+-
An aqueous kava rhizome extract, kawain, dihy- evoked contractions of skinned muscles. The
drokawain, methysticin and dihydromethysticin results suggested that the kava pyrone (±)-kavain
inhibited serotonin and nicotine-induced contrac- may act in a non-specific musculotropic way on
tions of guinea-pig ileum in-vitro (Meyer 1965; the smooth muscle membrane. In isolated iso-
Kretzschmar et al. 1969). The antispasmodic metrically contracted murine tracheal ring prepa-
effects were attributed to a direct musculotropic rations, kavain was observed to diminish the
action. Dihydromethysticin also inhibited con- maximal contractile response to both muscarinic
tractions of rat colon and uterus in-vitro induced receptor activation and voltage-operated calcium
by serotonin, acetylcholine and barium (Meyer channel activation (Martin et al. 2000). The IC50
1965). Desmethoxyyangonin, dihydromethysti- for kavain in rings precontracted with carbachol
cin and kawain inhibited serotonin-induced con- was found to be 177 μM, and, in rings precon-
tractions of rat uterus in-vitro at concentrations tracted with KCl, it was found to be 59.6 μM. In
of 3.2, 7.5 and 10.0 μg/ml, respectively (Buckley addition, pretreatment with kavain attenuated air-
et al. 1967). Aqueous, dichloromethane and way smooth muscle contraction evoked with
lyophilized extracts of the rhizome induced either carbachol or KCl. The EC50 for KCl was
relaxation of rat uterus in-vitro (ED50 22.5 μg/ml) not affected by kavain pretreatment. However,
(O’Hara et al. 1965). Singh (1983) investigated the EC50 for carbachol was significantly affected
the effects of an aqueous extract of kava rhizome by a high kavain pretreatment dose. Further,
on muscle contractility and neuromuscular trans- Martin et al. (2002) reported that kavain (10−6 M
mission in mouse hemidiaphragms and frog sar- to 10−3 M) relaxed rat aortic rings precontracted
torius muscles in-vitro using twitch tension and with phenylephrine (PE) in a dose-dependent
intracellular recording techniques. Kava extract manner. This response was not dependent on
(2–5 mg/ml) induced muscle relaxation by direct functional endothelium. In addition, kavain pre-
action on muscle contractility rather than by inhi- treatment (3 × 10−5 M or 3 × 10−4 M) attenuated
bition of neuromuscular transmission. It was con- vascular smooth muscle contraction evoked by
cluded that kava caused paralysis by mechanisms PE. However, kavain failed to attenuate
similar to local anaesthetics. PE-mediated contraction in calcium (Ca2+)-free
(±)-Kavain (1 μM/mM) dose-dependently buffer, indicating that intracellular signaling pro-
reduced contractions of isolated guinea-pig ileum cesses were likely not affected. Also, kavain did
evoked by carbachol (10 μM), by BAY K 8644 attenuate the contraction elicited by the adminis-
(0.3 μM) or by substance P (0.05 μM) (Seitz et al. tration of Ca2+ to depolarized tissue. Also, in
1997a). (±)-Kavain also inhibited the contractile rings pre-treated with the selective l-type Ca2+
responses induced by raising the extracellular K+ channel blocker nifedipine, kavain-mediated
concentration from 4 to 20 mM and by blocking relaxation was inhibited. Lastly, in rings
170 Piperaceae

selectively contracted with an l-type calcium kava pyrones, desmethoxyyangonin and

channel activator, kavain elicited dose-dependent (±±)-methysticin, displayed a competetive inhi-
(and ultimately complete) relaxation. These data bition pattern with mean Ki 0.28 μoM and
strongly suggested that kavain attenuated vascu- 1.14 μM, respectively. (+)-Kavain, a 4-methoxy-
lar smooth muscle contraction, likely through α-pyrone prepared from was found to have anti-
inhibition of Ca2+ channels. thrombotic action on human platelets as
evidenced from its ability to suppress arachidonic
acid (AA)-induced aggregation, exocytosis of
Antitrypanosomal Activity ATP and inhibition of cyclooxygenase (COX)
and thromboxane synthase (TXS) activity, the
Otoguro et al. (2012) found two phenolic com- latter two effects being estimated from the gen-
pounds β-phenethyl caffeate, farnesyl caffeate eration of prostaglandin E2 (PGE2) and throm-
from propolis and three kawa lactones kawain, boxane A2 (TXA2), respectively (Gleitz et al.
dihydrokawain and yangonin possessing an 1997). An application of (+)-kavain 5 min before
α-pyrone influenced antitrypanosomal property AA, dose-dependently diminished aggregation,
against Trypanosoma brucei brucei. In particular, ATP-release and the synthesis of TXA2 and PGE2
β-phenethyl caffeate, farnesyl caffeate and dihy- with IC50 values of 78, 115, 71 and 86 μmol/l,
drokawain exhibited high or moderate selective respectively.
and potent antitrypanosomal activity in-vitro.

Antimicrobial Activity
Melanogenesis Stimulation Activity
Kavain was reported to exhibit bactericidal activ-
P. methysticum rhizome ethanol extract showed ity against Gonococcus, the pathogenic agent of
potent stimulatory effect on melanogenesis in gonorrhoea and colon bacillus and blennorrhea
cultured murine B16 melanoma cells (Matsuda (Steinmetz 1960). The pathogenic fungi
et al. 2006). Activity-guided fractionation of Trichophyton ferrugineum, Trichophyton
kava extract led to the isolation of two active kav- tonsurans and Cryptococcus neoformans were
alactones, yangonin and 7,8-epoxyyangonin, strongly inhibited by kava extracts (Hänsel et al.
along with three inactive kavalactones, 1966a). Dihydromethysticin completely inhib-
5,6-dehydrokawain, (+)-kawain and (+)-methys- ited the growth of Aspergillus niger in-vitro
ticin, and a glucosylsterol, daucosterin. (Shulgin 1973). The fungistatic principles of the
7,8-Epoxyyangonin showed a significant stimu- kava root were found to be 4-methoxy-α-pyrones
latory effect on melanogenesis in B16 melanoma like dihydrokawain which completely inhibited
cells. Yangonin exhibited a weak melanogenesis Aspergillus niger at a concentration of 0.5 mg/
stimulation activity. ml. Other fungi (species of Candida and
Fusarium) and bacteria were not inhibited by
kava and dihydrokawain. A hydroalcoholic
Antiplatelet Activity extract of kava rhizome inhibited the growth in-
vitro of Aspergillus fumigatus, A. niger,
Kava-kava extract was found to be a reversible Penicillium digitatum, Rhizopus nigricans,
inhibitor of MAO-B in intact platelets (IC50 Trichophyton mentagrophytes, Candida albicans
24 μM) and disrupted platelet homogenates (IC50 and Saccharomyces pastorianus (Guérin and
1.2 μM) (Uebelhack et al. 1998). The order of Réveillère 1984). However, an aqueous extract of
potency was desmethoxyyangonin > (±)-methys- the rhizome weakly inhibited the growth in-vitro
ticin > yangonin > (±)-dihydromethysticin > (±)- of Trichophyton rubrum, Microsporum canis or
dihydrokavain > (±)-kavain. The two most potent Epidermophyton floccosum (Locher et al. 1995).
Piper methysticum 171

Amoebicidal Activity cytochrome P450 (P450) enzymes activities, with

significant inhibition of the activities of CYP1A2
The amoebicidal activity of yangonin was found (56 % inhibition), 2C9 (92 %), 2C19 (86 %), 2D6
comparable to that of the commercial drugs (73 %), 3A4 (78 %) and 4A9/11 (65 %) following
enterovioform, enteroquinol, clefamide and preincubation for 15 min with human liver micro-
furamidazole (Hänsel 1968; Sotheeswaran 1987). somes (HLMs) and NADPH; CYP2A6, 2C8 and
2E1 activities were unaffected (Mathews et al.
2002). Of its major kavalactones, kavain did not
Kava-Drug Interaction Activity inhibit these enzymes, there was significant inhi-
bition of CYP2C9 by desmethoxyyangonin
Kava (Piper methysticum) was one of the herbal (42 %), methysticin (58 %) and dihydromethysti-
remedies listed in the United States with the cin (69 %); 2C19 by dihydromethysticin (76 %);
potential to significantly modulate the activity of 2D6 by methysticin (44 %); and 3A4 by desme-
drug-metabolizing enzymes (notably cytochrome thoxyyangonin (40 %), methysticin (27 %) and
P450 isozymes) and/or the drug transporter dihydromethysticin (54 %). These data indicated
P-glycoprotein and could have potential adverse that kava possessed a high potential for causing
interactions with anticancer agents (Sparreboom drug interactions through inhibition of P450
et al. 2004). enzymes responsible for the majority of the
Studies had shown that several kavalactones, metabolism of pharmaceutical agents. Effects
the active principles of kava extracts, to be potent reported for the kava extracts may result from the
inhibitors of several enzymes of the CYP 450 different preparation protocols used. In cDNA-
system (CYP1A2, 2C9, 2C19, 2D6, 3A4 and expressed human enzymes and cryopreserved
4A9/11) indicating that kava had a high potential human hepatocytes, the kava extract and the three
for causing pharmacokinetic drug interactions kava lactones (methysticin, desmethoxyyangonin
with other herbal products or drugs, which are and yangonin) were found to be potent inhibitors
metabolised by the CYP 450 enzymes (Anke and of a panel of P450 isoforms (CYPs 1A2, 2C9,
Ramzan 2004). Analysis of a kava kava and a 2C19, 2E1 and 3A4) with IC50 values of approxi-
basil extract showed that the on-line HPLC sys- mately 10 μM (Zou et al. 2004b). The test com-
tem was applicable to complex mixtures, since in pounds were also moderately cytotoxic to human
both extracts, peaks with human cytochrom P450 hepatocytes (EC50 values of approximately
1A2 inhibiting activity were observed (Jeurissen 50 μM). Methysticin was the most potent enzyme
et al. 2007). The results of studies by Guo et al. inhibitor as well as the most cytotoxic, followed
(2009) indicated that kava extract could signifi- by (in order of potency) the kava root extract, des-
cantly modulate drug metabolizing enzymes, methoxyyangonin and yangonin. Commercial
particularly the CYP isozymes, which could kava extracts (prepared in acetone, ethanol or
cause herb-drug interactions and may potentially methanol) exhibited more pronounced inhibition
lead to hepatotoxicity. of P450 enzymes (CYP3A4, CYP1A2, CYP2C9
Almeida and Grimsley (1996) reported a case and CYP2C19) than traditional kava (aqueous)
of a 54-year-old man hospitalised in lethargic and extract (Côté et al. 2004). Among kava kavalac-
disoriented state and found to be induced by pos- tones, only desmethoxyyangonin and dihydrome-
sible drug interaction between kava a-pyrones and thysticin markedly induced the expression of
a benzodiazepine (alprazolam). Inhibitors of cyto- CYP3A23 (approximately 7-fold) in rat hepato-
chrome P450 3A4 (CYP3A4) were identified in cytes (Ma et al. 2004). The results suggested that
crude extracts from kava roots (Unger et al. 2002). induction of CYP3A23 by dihydromethysticin
Kavapyrones were identified as the main CYP3A4 and desmethoxyyangonin involved transcription
inhibitory components of kava. Whole kava activation, probably through a human pregnane X
extract (normalized to 100 μM total kavalactones) receptor (PXR)-independent or PXR-involved
caused concentration-dependent decreases in indirect mechanism.
172 Piperaceae

Treatment of F-344 rats with pipermethystine methysticin triggered the most profound induc-
(10 mg/kg) and kava root acetone-water extract ing effect on CYP1A1 followed by 7,8-dihydro-
(100 mg/kg) for 2 weeks failed to elicit any sig- methysticin. The other four kavalactones
nificant changes in liver function tests or cause (yangonin, 5,6-dehydrokawain, kawain and
severe hepatic toxicity as measured by lipid per- 7,8-dihydrokawain) did not show significant
oxidation and apoptosis markers such as malo- effects on CYP1A1. It was found that kava
ndialdehyde, Bax and Bcl-2 (Lim et al. 2007). extract induced the expression of CYP1A1 via an
However, pipermethystine-treated rats demon- aryl hydrocarbon receptor (AhR)-dependent
strated a significant increase in hepatic glutathi- mechanism and that methysticin and
one, cytosolic superoxide dismutase (Cu/ 7,8-dihydromethysticin contributed to CYP1A1
ZnSOD), tumor necrosis factor alpha mRNA induction. The induction of CYP1A1 indicated a
expression and cytochrome P450 (CYP) 2E1 and potential interaction between kava or kavalactones
1A2, suggesting adaptation to oxidative stress and CYP1A1-mediated chemical carcinogenesis.
and possible drug-drug interactions. Yamazaki Weiss et al. (2005) demonstrated P-glycoprotein
et al. (2008) found that a high dose (equivalent to inhibitory activity of kava-kava and its
approximately 380 mg kavalactones/kg/day; 100 kavalactones kavalactones kavain, dihydrokavain,
times of the suggested dosage for human use) of methysticin, dihydromethysticin, yangonin and
two different types of kava products for 8 days desmethoxyyangonin in P-glycoprotein-over-
significantly increased liver weights in rats. expressing cell line P388/dx and the correspond-
CYP1A2 mRNA expression was moderately ing cell line P388. The crude extract and the
increased (2.8–7.3 fold). More importantly, the kavalactones showed a moderate to potent inhibi-
high dose of kava markedly enhanced CYP1A1 tory activity with f2 (concentration needed to
mRNA expression (75–220 fold) as well as double baseline fluorescence) values of 170 μg/ml
ethoxyresorufin O-deethylase activities and and 17–90 μM, respectively. The f2 value of yan-
CYP1A1 immunoreactivities. Thus, no observed gonin could not be determined due to its higher
adverse effect levels of kavalactones would be lipophilicity. Mathews et al. (2005) found that
lower than 380 mg/kg/day. They advised that kava could cause adverse drug reactions via inhi-
considerable attention should be paid to the pos- bition of drug metabolism. The 7-day pretreat-
sibility that kava products could induce hepatic ment with kava extract only modestly induced
CYP1A1 expression in human especially in sen- hepatic cytochrome P450 activities. The human
sitive individuals. hepatic microsomal P450s most strongly inhib-
Gene expression profiling in male B6C3F1 ited by kava extract (CYP2C9, CYP2C19,
mouse livers administered kava extract by gavage CYP2D6, CYP3A4) were inhibited to the same
for 14 weeks identified the differentially degree by a “composite” kava formulation com-
expressed drug metabolizing genes in response to posed of the six major kavalactones contained in
kava treatments (Guo et al. 2010). It was found the extract. Ki values for the inhibition of CYP2C9
that the levels of significant numbers of genes and CYP2C19 activities by methysticin, dihydro-
involving drug metabolism were changed and methysticin and desmethoxyyangonin ranged
that the pathways involving xenobiotics metabo- from 5 to 10 μM. Kava extract and kavalactones
lism, Nrf2-mediated oxidative stress response, (< or =9 μM) modestly stimulated P-glycoprotein
mitochondrial functions and others were altered. ATPase activities. Two cases of patients seen on
Their results indicated that kava extract can the psychiatric emergency and consult service
significantly modulate drug metabolizing who developed severe side effects from psycho-
enzymes, potentially leading to herb-drug inter- tropic medications in the context of kava use were
actions and hepatotoxicity. Studies showed that reported by Toohey et al. (2013). In both cases,
kava extract displayed a concentration-dependent kava use may have affected the metabolism of the
effect on hepatic CYP1A1 induction (Li et al. psychotropic medications, leading to serious side
2011). Among the six major kavalactones, effects. Growing research indicated that kava
Piper methysticum 173

most likely altered concentrations of co-adminis- vidual injection. Dihydrokawain and desme-
tered psychotropics possibly by inhibiting cyto- thoxyyangonin levels remained the same as those
chrome P450 enzymes. established for their individual administration.
Methane chemical ionization (CI) gas chroma-
tography–mass spectrometry (GC–MS) was used
Pharmacokinetic Studies by Duffield et al. (1989a) to identify some of the
human urinary metabolites of the kava lactones
After the administration of kava pyrones to rats, following the ingestion of aqueous kava extract.
about half the dose (400 mg/kg, p.o.) of dihydro- All seven major, and several minor, kava lactones
kawain was found in the urine in 48 h, about two- were identified in human urine. Observed meta-
thirds of this was hydroxylated metabolites (three bolic transformations include the reduction of the
mono- and three di-hydroxylated derivatives), of 3,4-double bond and/or demethylation of the
which p-hydroxydihydrokawain was the most 4-methoxyl group of the α-pyrone ring system.
abundant (Rasmussen et al. 1979). The remain- Demethylation of the 12-methoxy substituent in
ing third consisted of metabolites formed by yangonin (or alternatively hydroxylation at C-12
scission of the 5,6-dihydro-alpha-pyrone ring of desmethoxyyangonin) was also observed. In
and included hippuric acid (9–13 % dose). contrast to the situation prevailing in the rat, no
Lower amounts of urinary metabolites were dihydroxylated metabolites of the kava lactones,
excreted when kawain was administered, but or products from ring opening of the 2-pyrone
both hydroxylated and ring-opened products ring system, were identified in human urine.
were formed. Methysticin afforded only small Abourashed and Khan (2000) used fungal models
amounts of two urinary metabolites formed by to mimic the mammalian metabolism of two major
demethylenation of the methylenedioxyphenyl kava styryl alpha-pyrones, d-kawain and d-methys-
moiety. Urinary metabolites of the alpha-pyrones, ticin. The culture broth of Cunninghamella elegans
7,8-dihydroxyangonin and yangonin were yielded 4′-hydroxykawain from d-kawain, the
formed via omicron-demethylation. No ring- same metabolite identified in rat urine. The fungus
opened products were detected. These lipophilic Torulopsis petrophilum biotransformed d-methys-
kava pyrones had extremely low solubility in ticin to 3′-hydroxy-4′-methoxykawain which was
water, which would be expected to reduce their analogous, but not identical, to a known rat metab-
absorption rates and appeared to be responsible olite of methysticin. Kavalactones appeared to be
for the variable and low extent of metabolism hydroxylated by the cytochrome P450 system and
observed. their metabolism may be enhanced by the presence
Intraperitoneal (i.p.) administration of mice of glutathione (Russmann et al. 2001; Tinsley
with 100 mg/kg of dihydrokawain, kawain, des- 1999).
methoxyyangonin and yangonin was investigated The oral pharmacokinetics of the kavalactone,
in relation to uptake by mouse brain (Keledjian kawain (100 mg/kg), were determined in rats
et al. 1988). After 5 min, dihydrokawain and with and without coadministration of kava extract
kawain attained maximum concentrations of 64.7 (256 mg/kg) to study the effect of the extract on
and 29.3 ng/mg wet brain tissue, respectively. drug disposition (Mathews et al. 2005). Kawain
Desmethoxyyangonin and yangonin had poorly was well absorbed, with >90 % of the dose elimi-
defined maxima corresponding to concentrations nated within 72 h, chiefly in urine. Compared
of 10.4 and 1.2 ng/mg wet brain tissue, respec- with kawain alone, co-administration with kava
tively, and these compounds were more slowly extract caused a tripling of kawain AUC (0–8 h)
eliminated from brain tissue. When crude kava and a doubling of Cmax. However, a 7-day pre-
resin was administered i.p. at a dose of 120 mg/ treatment with kava extract (256 mg /kg/day) had
kg, the concentrations of kawain and yangonin no effect on the pharmacokinetics of kawain
markedly increased (2 and 20 times, respectively) administered on day 8. Studies by Fu et al. (2012)
relative to the values measured from their indi- found among the six major kavalactones (kavain,
174 Piperaceae

methysticin and desmethoxyyangonin) and high-dose dogs, Kavaform was mildly toxic; pro-
their respective metabolites (p-hydroxykavain, liferation of the small cells of the thyroid epithe-
m,p-dihydroxykavain and p-hydroxy-5,6- lium and a multicentric necrosis of the
dehydrokavain), p-hydroxykavain and m,p-di- parenchyma of the liver were observed as a single
hydroxykavain were the only metabolites histological finding in one high-dose dog. Hsu
detected in the perfusate of rat liver. Kavalactone et al. (1994) tested dihydrokavain and desme-
biliary excretion was negligible. Kavalactones thoxyyangonin in mice and Wistar rats on oral
were found to be potentially bioavailable as they application. An application of 30, 100 or 300 mg/
all readily crossed the Caco-2 cell monolayers kg twice a day for 2 weeks did not yield any clini-
with apparent permeabilities (Papp) increasing cally relevant changes in haematological and his-
from 42 × 10−6 cm/s and most exhibiting more tological parameters. Rats were administered
than 70 % crossing within 90 min (Matthias et al. similar dosages over 3 months. Only with the use
2007). Not all differences in their bioavailability of desmethoxyangonin, transient changes of
can be related to kavalactone structural differ- serum glucose, cholesterol and triglycerides were
ences as it appeared that bioavailability may also found, however, without a clear dosage–effect
be affected by co-extracted compounds. For relationship. Overall, the experiment did not
example, the Papp for kawain from ethanol extracts yield hints on toxic effects of kavalactones.
was higher than the values obtained for the same Feeding Wistar rats of both sexes with 7.3 or
compound from water extracts or for the kavalac- 73 mg/kg body weight of ethanolic kava extract
tone alone. The pilot study by Tarbah et al. (2013) for 3 and 6 months elicited no signs of toxicity in
found that kavalactones (kavain; 7,8-dihydrokavain, relation to changes in body weight, hematologi-
methysticin; 7,8-dihydromethysticin; 5,6-dehydro- cal and liver parameters, and macroscopical and
kavain (=desmethyoxyyangonin); and yangonin) microscopical histological changes in the major
accumulated in the keratin matrix of hair and could organs (Sorrentino et al. 2006).
provide an easily applicable system for assessing None of the six major kavalactones was found
chronic consumption of kava. to be positive in the experimental concentration
ranges tested by the umu test (a sensitive test for
point mutations) (Jhoo et al. 2007). However,
Toxicological Studies among the different solvent fractions, the
n-butanol fraction of kava leaves was positive for
Oral administration of a test mixture of 40 % mutagenicity. Further investigations using
kavain, 40 % dihydrokavain, and 20 % yangonin bioassay-directed isolation and analysis indicated
at doses of 100 or 500 mg/kg on days 6–15 of that 2 C-glycoside flavonoid compounds identi-
gestation.was found not teratogenic or embryo- fied as 2″-O-rhamnosylvitexin and schaftoside
toxic in Wistar rats (Hapke et al. 1971). The mix- accounted for the positive mutagenic results.
ture was also negative for teratogenic activity in Whittaker et al. (2008) evaluated the toxicity and
New Zealand strain rabbits when administered mutagenicity of two commercial samples of
orally at doses of 20 or 200 mg/kg on days 6–18 kava, Kaviar and KavaPure, and the six pure kav-
after mating. However, there was a significant alactones, kawain, dihydrokawain, methysticin,
dose-related reduction of foetal weight in treated dihydromethysticin, yangonin and desmethoxyy-
rabbits. In subchronic toxicological studies of angoni including both d-kawain and dl-kawain,
Kavaform®, a German geriatric preparation con- in L5178Y mouse lymphoma cells. They found
taining 50 mg of d,1-kavain and 200 mg of mag- neither the kava samples nor the kavalactones
nesium orotatein in Wistar rats and mongrel induced a mutagenic response in the L5178Y
dogs, serum glutamate pyruvate transaminase mouse lymphoma mutation assay with the addi-
(SGPT) levels were significantly increased in tion of human liver S9 activation.
high-dose rats; however, liver cell damage was Under the conditions of the 2-year toxicology
not confirmed by histological examination. In and carcinogenesis studies of kava kava extract
Piper methysticum 175

(CAS No. 9000-38-8) in F344/N rats and B6C3F1 severe hepatotoxic reactions to kava. Incubation
mice (gavage studies) by US Department of of kava lactones (methysticin, yangonin and des-
Health and Human Services, National Toxicology methoxyyangonin) and an ethanolic extract of
Program, there was equivocal evidence of carci- dried kava root in culture with MCL-5 cells, a
nogenic activity of kava kava extract in male human lymphoblastoid cell line stably trans-
F344/N rats based on marginal increases in the fected with five human P450’s (CYP 1A1, 1A2,
incidences of testicular interstitial cell adenoma 2A6, 2E1 and 3A4) and human epoxide hydro-
(NTP 2012). There was no evidence of carcino- lase and a control cell line (cH2) derived from the
genic activity of kava kava extract in female same parental line as MCL-5, but transfected
F344/N rats administered 0.1, 0.3 or 1.0 g/kg. with two empty vectors, elicited varying degrees
There was clear evidence of carcinogenic activity of metabolic toxicity (IC50 values ranged from
of kava kava extract in male B6C3F1 mice based 50 to > 100 μM) to both MCL-5 and cH2 cell
on increased incidences of hepatoblastoma. lines; however, both cell lines were equally sensi-
There was some evidence of carcinogenic activ- tive to the test compounds (Zou et al. 2004a). The
ity of kava kava extract in female B6C3F1 mice results suggested that the parent compound for
based on increased incidences of hepatocellular each of the four test compounds was primarily
adenoma or carcinoma (combined). In gentic responsible for the observed cell toxicity and that
toxicology, kava kava extract was negative in CYP 1A1, 1A2, 2A6, 2E1 and 3A4 or epoxide
Salmonella typhimurium strains TA97, TA98, hydroxylase did not appear to be involved. It was
TA100 and TA1535 with and without S9 activa- concluded that in-vitro kava did not appear to be
tion and also negative in Escherichia coli WP2 activated to toxic metabolites by enzymes known
uvrA/pKM101 with and without S9. In-vivo, no to be important in metabolic toxicity. Studies by
increases in the frequencies of micronucleated Whitton et al. (2003) found that the extraction
erythrocytes were observed in peripheral blood process (aqueous vs. acetone in the two types of
of male or female B6C3F1 mice administered preparations) was responsible for the difference
kava kava extract by gavage for 3 months. in toxicity as extraction of glutathione in addition
to the kava lactones was important to provide
protection against hepatotoxicity. The Michael
Kava Hepatotoxicity reaction between glutathione and kava lactones,
resulting in opening of the lactone ring, reduced
In-Vitro Studies the side effects of the kava kava extracts. This
Exposure of human hepatoma cells, HepG2, to protective activity was demonstrated using
100 μM pipermethystine caused 90 % loss in cell Acanthamoebae castellanii in which 100 % cell
viability within 24 h, while 50 μM caused 65 % death occurred with 100 mg/ml kava lactones
cell death (Nerurkar et al. 2004). Similar concen- alone, and 40 % cell death with a mixture of
trations of kavalactones (7,8-dihydromethysticin 100 mg/ml glutathione and 100 mg/ml kava
and desmethoxyyangonin) did not affect cell via- lactones.
bility for up to 8 days of treatment. Mechanistic Jhoo et al. (2006) found that organic solvent
studies indicated that, in contrast to kavalactones, fractions displayed a much stronger cytotoxicity
pipermethystine significantly decreased cellular than water fractions for kava root, leaf and
ATP levels and mitochondrial membrane poten- stem peelings as evaluated in HepG2 cells
tial and induced apoptosis as measured by the based on the 3-(4,5-dimethylthiazol-2-yl)-2,5-
release of caspase-3 after 24 h of treatment. diphenyltetrazolium bromide assay and lactate
These observations suggested that pipermethys- dehydrogenase and aspartate aminotransferase
tine, rather than kavalactones, was capable of enzyme leakage assays. The hexane fraction of
causing cell death, probably in part by disrupting the root exhibited stronger cytotoxic effects than
mitochondrial function. Thus, it was concluded fractions of root extracted with other solvents
that pipermethystine may contribute to rare but (ethyl acetate, n-butanol and water) or extracts
176 Piperaceae

from the other parts of kava. Further investiga- root, to be a potent hepatocellular toxin, inducing
tions using bioassay-directed isolation and analy- cell death in HepG2 (LD50 = 15.3 μM) and l-02
sis of the hexane fraction indicated that the (LD50 = 32 μM) cells. Hepatocellular toxicity of
compound responsible for the cytotoxicity was FKB was mediated by induction of oxidative
flavokavain B. stress, depletion of reduced glutathione (GSH),
The pyridone alkaloid pipermethystine had inhibition of IKK activity leading to NF-κB tran-
been considered to be responsible for alleged scriptional blockade and constitutive TNF-α-
hepatoxicity of Kava products (Lechtenberg et al. independent activation of mitogen-activated
2008). Investigation of a series of retain samples protein kinase (MAPK) signaling pathways,
of finished products from the German market and namely, ERK, p38 and JNK. They further dem-
self-produced extracts from root and stem mate- onstrated by non-invasive bioluminescence imag-
rial of Piper methysticum clearly showed that ing that oral consumption of FKB leads to the
pipermethystine was absent from all root and inhibition of hepatic NF-κB transcriptional activ-
retain samples and extracts, with a limit of quan- ity in-vivo and severe liver damage.
tification of 45 ppm. Thus they stated that piper- Hepatotoxicity could occur as an acute, severe
methystine should not be the responsible form or a chronic, mild form in relation to kava
constituent in hepatoxic case reports with ethano- ingestion (Rowe et al. 2011). Inflammation
lic kava extracts produced in Germany. appeared to be involved in both forms and may
Studies by Lüde et al. (2008) found kava result from the activation of liver macrophages
extracts were toxic to mitochondria, leading to (Kupffer cells), either directly or via kava metab-
inhibition of the respiratory chain, increase in olites. Pharmacogenomics may influence the
ROS production, decrease in the mitochondrial severity of this inflammatory response. According
membrane potential and eventually to apoptosis to Clouatre (2004), three possible mechanisms
of exposed HepG2cells. The methanolic and an for kavalactone hepatotoxicity were known: inhi-
acetonic kava root and a methanolic leaf extracts bition of cytochrome P450, reduction in liver glu-
showed cytotoxicity starting at a concentration of tathione content and, more remotely, inhibition
50 μg/ml (lactate dehydrogenase leakage) or of cyclooxygenase enzyme activity. The patho-
1 μg/ml (MTT test). The mitochondrial mem- physiology of kava hepatotoxicity remains
brane potential was decreased (root extracts start- inclonclusive despite the existence of circum-
ing at 50 μg/ml) and the respiratory chain stantial evidence for the roles of toxic metabo-
inhibited and uncoupled (root extracts) or only lites, inhibition of cyclooxygenase (COX)
uncoupled (leaf extract) at 150 μg/ml, and mito- enzymes and depletion of liver glutathione
chondrial beta-oxidation was inhibited by all (Zhang et al. 2011). Experimental and clinical
extracts starting at 100 μg/ml. Induction of apop- cases of hepatotoxicity showed evidence of
tosis was demonstrated by all extracts at a con- hepatitis.
centration of 150 μg/ml. They stated that in
predisposed patients, mitochondrial toxicity of Animal Studies
kava extract may explain hepatic adverse reac- Studies by Singh and Devkota (2003) found that
tions of this drug. Kava was found to potentiate aqueous kava extract did not affect liver function
acetaminophen-induced hepatocyte cytotoxicity in rats. The data showed that none of the enzymes
(Yang and Salminen 2011). The findings indi- such as alanine aminotransferase, aspartate ami-
cated that kava potentiated acetaminophen- notransferase, alkaline phosphatase,lactate dehy-
induced cytotoxicity of rat primary hepatocytes drogenase andmalondialdehyde were elevated; in
by increasing the magnitude of glutathione deple- fact in some cases they were significantly
tion, resulting in oxidative stress and mitochon- reduced, suggesting the lack of a toxic effect by
drial dysfunction, ultimately leading to cell death. kava on the liver. Separate studies by DiSilvestro
Studies by Zhou et al. (2010) demonstrated et al. (2007) found 3 months of kava feeding in
that flavokawain B (FKB), a chalcone from kava rats at three different doses (31.25, 62.5 and
Piper methysticum 177

133 mg/kg diet) produced no liver injury based study of human cancers. Zhang et al. (2013)
on serum markers of liver damage (sorbitol dehy- found that liver macrophage depletion amelio-
drogenase activities, bile acid concentrations and rated kavalactone damage in isolated perfused rat
beta-glucuronidase activities) and serum lipid liver.
peroxide readings. For some measurements and In prechronic studies, orally administered
some kava doses, the injury marker readings kava at 0.125–2 g/kg body weight revealed dose-
were below control values. Moreover, for these related increases in liver weights and incidences
same parameters, kava feeding did not enhance of hepatocellular hypertrophy in F344/N rats and
the effects of the hepatotoxin galactosamine B6C3F1 mice (Behl et al. 2011). In the chronic
(500 mg/kg ip); some kava doses even showed studies, there were dose-related increases in the
modest protection against liver injury. Liver his- incidences of hepatocellular hypertrophy in rats
tology analysis showed no signs of kava causing and mice administered kava kava for up to 1 g/kg
or enhancing liver injury. Their study did not sup- body weight. This was accompanied by signifi-
port the concept that kava produces or aggravates cant increase in incidences of centrilobular fatty
liver injury. Studies by Narayanapillai et al. change. Male mice showed a significant dose-
(2014b) found that kava alone revealed no related increase in the incidence of hepatoblasto-
adverse hapatoxic effects for long-term usage mas. In female mice, there was a significant
even at a dose of 500 mg/kg bodyweight in increase in the combined incidence of hepatocel-
C57BL/6 mice. In contrast, a 3-day kava pre- lular adenoma and carcinoma in the low- and
treatment potentiated acetaminophen P-induced mid-dose groups but not in the high-dose group.
hepatotoxicity, resulted in an increased serum These findings were accompanied by several
ALT and AST and increased severity of liver nonneoplastic hepatic lesions.
lesions. It was found that flavokawains A and B
in kava, not dihydromethysticin, potentiated Clinical Studies/Reports
acetaminophen-induced hepatotoxicity. A case of a woman with acute necrotizing hepati-
Dietary feeding of flavokawain A (FKA), a tis after taking herbal remedies as alternative
major kava chalcone, did not affect food con- medication, containing kava, was reported in
sumption and body weight of male FVB/N mice Germany in 1998 (Strahl et al. 1998). Viral, auto-
(Li et al. 2014). Histopathological examination immune and metabolic causes of the hepatitis
of liver, kidney, colon, lung, heart, spleen and were excluded. Campo et al. (2002) reported a
thymus revealed no signs of FKA-induced toxic- case of a girl aged 14 years who was hospitalised
ity. Biochemical serum analysis and histological with kava-induced fulminant hepatic failure in
examination confirmed normal organ function in December 2000, after taking two kava products
FKA-treated mice. The cytotoxicity profile from August to December. Initial liver biopsy
showed FKA had minimal side effects on bone revealed active fulminant hepatitis with extensive
marrow and small intestinal epithelial cells com- centrilobular necrosis, approximately 25 % hepa-
pared with adriamycin. In addition, oral feeding tocellular viability, and mixed inflammatory infil-
of FKA increased activities of both glutathione trates consisting of lymphocytes, histiocytes,
S-transferase and quinone reductase in the liver, scattered eosinophils and occasional neutrophils.
lung, prostate and bladder tissues of mice. In No viral cytopathic changes were identified, and
comparison, dietary feeding of 0.6 % commercial immunohistochemical stains for hepatitis B sur-
kava root extract increased liver/body weight face and core antigens were negative. Humberston
ratio and decreased spleen, thymus and testis/ et al. (2003) described a case of a previously
body weight ratios, as well as induced nodular healthy 14-year-old female was admitted to the
proliferation in liver tissues. Therefore, dietary hospital with hepatic failure. Initial therapy,
feeding FKA showed no adverse effects on major including plasmapheresis, was unsuccessful and
organ function and homeostasis in mice, suggest- she deteriorated. She ultimately required a liver
ing the potential of FKA for chemoprevention transplant and now remains well. The liver biopsy
178 Piperaceae

showed hepatocellular necrosis consistent with sionflower (Passiflora incarnata). The patient
chemical hepatitis. A work-up for alternative died after a report by the Therapeutic Goods
causes of liver failure was negative. The patient Administration (TGA) warning of the potential
gave a history of taking a kava kava – containing for hepatotoxicity associated with the use of
product for 4 months. kava-containing products. In a study of 31 healthy
Russmann et al. (2003) reported that tradi- adult Tongan kava drinkers versus a control group
tional aqueous kava extracts were the most prob- of 31 healthy adult Tongan non-kava drinkers in
able cause of hepatitis in two patients presenting Oahu Hawaii, it was found that chronic heavy
with markedly elevated transaminases and hyper- kava beverage consumption was associated with
bilirubinaemia. A consequent survey of 27 heavy the elevation of γ-glutamyl transpeptidase in
kava drinkers in New Caledonia showed elevated 65 % of the kava drinkers versus 26 % in the con-
gamma glutamyl transferase in 23/27 and mini- trols (Brown et al. 2007). Alkaline phosphatase
mally elevated transaminases in 8/27. They con- was elevated in 23 % of kava drinkers versus 3 %
cluded that not only commercially available but in the controls.
also traditionally prepared kava extracts may In an otherwise healthy 48-year-old female
rarely cause liver injury. The increased activity of patient, acute hepatitis with transaminase increase
gamma glutamyl transferase in heavy kava con- (glutamic oxaloacetic transaminase (GOT) up to
sumers in the presence of normal or minimally 613 U/l, glutamic pyruvic transminase (GPT) up
elevated transaminases was probably not a sign to 752 U/l), inconspicuous hepatitis serology
of liver injury, but rather reflected an induction of findings, negative autoantibody status and nega-
CYP450 enzymes. In a cross-sectional study with tive virus serology was observed after a 10-week
98 indigenous Australian participants, 36 of long intake of kava kava (1–3 × 200 mg/day) and
whom had never used kava, it was found that St John’s Wort (1 × 425 mg/day) (Musch et al.
more recent kava use was independently associ- 2006). Biopsy of the liver showed lobular and
ated with higher levels of liver enzymes gamma- portal necroinflammatory activity without indica-
glutamyl transferase (GGT) and alkaline tion of cirrhosis. Discontinuation of the existing
phosphatase (ALP), but not with alanine amino- medication and simultaneous onset of immuno-
transferase or bilirubin, which were not elevated suppressive combination therapy of cortisone,
(Clough et al. 2003a). In those who were not azathioprine and ursodeoxycholic acid resulted
heavy alcohol users, only those who used kava in normalisation of the liver parameters within a
within the previous 24 h showed GGT levels period of 2 months. It appeared that simultaneous
higher than nonusers, whereas higher ALP levels intake of St John’s Wort possibly potentiated the
occurred only in those who last used kava 1–2 toxicity of kavapyrones. On the other hand, an
weeks and 24 h previously. It was concluded that immune-mediated mechanism, induced by kava
liver function changes in users of aqueous kava kava, could not be completely excluded. A com-
extracts at these moderate levels of consumption parative study of two structured quantitative ana-
appeared to be reversible and may begin to return lytical methods: the system of Maria and
to baseline after 1–2 weeks abstinence from kava. Victorino (MV) and that of the Council for
No evidence for irreversible liver damage had International Organizations of Medical Sciences
been found. In another paper, Clough et al. (CIOMS) for causality assessment of kava hepa-
(2003b) conducted a cross-sectional study of 101 totoxicity was carried out on 26 patients with
aboriginal adults in east Arnhem Land in the assumed hepatotoxicity by the herb (Teschke
Northern Territory. They found that kava use was et al. 2010). Grades of causality for suspected
associated with dermopathy, liver function abnor- hepatotoxicity by kava were much lower when
malities and decreased lymphocytes. Gow et al. evaluated by structured quantitative causality
(2003) described a case of acute liver failure and assessment scales than by regulatory ad hoc
death associated with the use of a preparation judgements. The quantitative CIOMS scale was
containing the “natural” anxiolytic kava and pas- found to be the preferable tool for causality
Piper methysticum 179

assessment of spontaneous reports of hepatotoxcity of 26 suspected cases of kava hepatotoxicity,

involving kava. Teschke et al. (2008) found that kava taken as
recommended to be associated with rare hepato-
Review/Analytical Studies toxicity, whereas overdose, prolonged treatment
Stickel et al. (2003) analyzed 29 novel cases of and co-medication may carry an increased risk.
hepatitis along with kava ingestion which In a comparative study with nine patients from
occurred between 1990 and 2002 in addition to Germany and Switzerland with established cau-
the seven already published case reports using a sality of hepatotoxicity by ethanolic and acetonic
clinical diagnostic scale established for adverse kava extracts and five patients from New
hepatic drug reactions. They found that hepatic Caledonia, Australia, the United States and
necrosis or cholestatic hepatitis were observed Germany for aqueous kava extracts and kava-
with both alcoholic and acetonic kava extracts. herbs mixtures, Teschke et al. (2009) in a com-
The majority of the 29 patients and the additional parative study of hepatotoxicty caused by
seven published reports were women (27 females, aqueous, acetonin and ethnaolic kava extracts
nine males). Both the cumulative dose and the found that the clinical picture in all 14 patients
latency to when the hepatotoxic reaction emerged was similar, independently whether aqueous,
were highly variable. Nine patients developed ethanolic and acetonic kava extracts or kava-
fulminant liver failure, of which eight patients herbs mixtures were used, substantiating that the
underwent liver transplantation. Three patients solvents employed to prepare the various kava
died, two following unsuccessful liver transplan- extracts were not causally related to the develop-
tation and one without. In all other patients, a ment of liver injury in these cases. This finding
complete recovery was noticed after the with- confirmed the WHO kava report (2007) that
drawal of kava. Their report highlighted the traditional aqueous kava extracts may exert rare
potentially severe hepatotoxicity of kava which potential hepatotoxicity similar to acetonic and
led to the retraction of kava-containing drugs by ethanolic extracts.
the pharmacovigilance authorities in Germany. In Teschke (2010) reviewed and discussed the
January 2003, kava extracts had been banned in possible pathogenetic factors for the develop-
the entire European Union and Canada, and were ment of kava-induced liver injury. He concluded
subject to cautions and advisories by the US FDA that kava hepatotoxicity occurred independently
as a result of 11 cases of hepatic failure leading to of the extraction medium used for the kava
liver transplants, including four deaths (Clouatre extracts and may primarily be attributed to daily
2004). A total of 78 cases of hepatotoxicity overdose, prolonged treatment and to a few kava
repeatedly linked to kava ingestion were avail- extract batches of poor quality; by improving
able for review from various databases. Teschke kava quality and adherence to therapy recom-
(2003) reported that almost 80 % of the patients mendation under avoidance of comedication,
took kavapyrones in overdose (maximally liver injury by kava should be a preventable dis-
480 mg/day) and/or for a prolonged time of more ease, at least to a major extent. Several reports of
than 3 months up to 2 years, which exceeded the hepatotoxicity had been linked to the consump-
commonly recommended daily dose of tion of kava extracts in Western countries, where
60–120 mg kavapyrones and the duration of the mainly ethanolic or acetonic extracts were used
therapy of up to 3 months for anxiety therapy. (Olsen et al. 2011). The mechanism of toxicity
Additional risks factors include co-medication had not been established, although several theo-
with up to five other chemically defined or herbal ries had proposed. On the basis of the chemical
drugs with in part potentially hepatotoxic proper- structures of kava constituents, the formation of
ties as well as a genetic deficiency of the hepatic reactive metabolites had also been suggested as a
microsomal cytochrome P450 2D6. Based on basis of toxicity. Furthermore, skin rash a side
findings of a clinical survey and critical analysis effect in kava consumers, may be indicative of
180 Piperaceae

the formation of reactive metabolites and cova- tion to kava hepatotoxicity, they (Teschke et al.
lent binding to skin proteins leading to immune- 2011d) proposed a six-point plan for new kava
mediated responses. Reactive metabolites of kava standardization: (1) use of a noble kava cultivar
lactones had been identified in-vitro as glutathi- such as Borogu, at least 5 years old at time of
one (GSH) conjugates and in-vivo as mercaptur- harvest; (2) use of peeled and dried rhizomes and
ates excreted in urine. Unfortunately, only a few roots; (3) aqueous extraction; (4) dosage recom-
studies had investigated the toxicity of the minor mendation of ≤ 250 mg kavalactones per day (for
constituents present in kava extract, such as medicinal use); (5) systematic rigorous future
pipermethystine and the flavokavains, where research; and (6) a Pan Pacific quality control
some have been shown to display higher in-vitro system enforced by strict policing. Teschke and
cytotoxicity than the lactones. They reiterated Lebot (2011) recommended for the establishment
that to date, there remains no indisputable reason of Pan-Pacific kava quality legislation as an
for the increased prevalence of kava-induced important part of the proposed Kava Quality
hepatotoxicity in Western countries. Teschke Standardization Code. They riterated that a
et al. (2012) in a review of cases analyzed by the sophisticated approach to establish kava quality
World Health Organization and published case standardizations was needed for safe human use
reports found that traditional aqueous extracts of kava as relaxing traditional beverages, anxio-
used in New Caledonia, Australia, the USA and lytic drugs and recreational dietary supplements.
Germany may also be hepatotoxic and may not Teschke et al. (2013) reported that kava hepato-
be restricted to use of Western acetonic and etha- toxicity was presently not reproducible experi-
nolic extracts. They stated that the primary cause mentally in preclinical models, as demonstrated
of toxicity may reside in the time before the prep- by studies showing whole kava extracts were not
aration of the various kava extracts, possibly hepatotoxic. This led them to propose the
attributed to poor quality of the raw material “working hypothesis” that contaminant hepato-
caused by mould hepatotoxins. They advised for toxins including moulds might have caused rare
rigorous testing of kava raw material, in addition kava hepatotoxicity in humans and that further
to Pan-Pacific kava manufacturing quality stan- studies were warranted to proof or disproof their
dards. According to Teschke et al. (2011a;b) thus working hypothesis. Currently, there is a paucity
far no clear evidence existed for the for a caus- of evidence supporting the recent speculation
ative hepatotoxic role of kavalactones and non- that adulterants or impurities such as the mould
kavalactone constituents, such as pipermethystine hepatotoxin aflatoxin could more likely cause
and flavokavain B identified from kava. They kava hepatotoxicity rather than kava components,
asserted that studies should focus on the identifi- such as kavalactones, pipermethystine and flavo-
cation of further potential hepatotoxic constitu- kavain B, that had demonstrated hepatotoxicity
ents, considering in particular possible kava (Rowe and Ramzan 2012). Although there was
adulterants and impurities with special reference limited similarity between acute kava hepatotox-
to ochratoxin A and aflatoxins (AFs) producing icity and acute aflatoxicosis, and background lev-
Aspergillus varieties. Also due to to high temper- els of aflatoxin had been detected in kava samples,
ature and humidity in the South Pacific area, kava there is an urgent need for epidemiological inves-
raw material might have been contaminated by tigations to uncover direct evidence implicating
mould hepatotoxins such as aflatoxins after har- mould hepatotoxins.
vest and during storage (Teschke et al. 2011b).
The analysis conducted by Teschke et al. (2011c)
showed poor quality of kava raw material as a Other Adverse Health/
cause for its liver toxicity and suggested preven- Dermatological Issues
tative measures by going back to the traditional
use of kava for the sake of the patients and the In a hospital-based health survey of 150 people
South Pacific economy. Consequently as a solu- on kava drinking in Vanuatu, Grace (2003) found
Piper methysticum 181

that 59 % of men and 15 % of women drank kava review of key evidence, their findings were as
(approximately 4:1). 51 % of all men drank kava follows: causality indicated: scaly skin rash,
at least weekly, compared to 11 % of women. For weight loss, raised gamma glutamyl transpepti-
any given kava-drinking episode, men drank dase liver enzyme levels, nausea, loss of appetite
more than women, (4.3 vs 3.3 shells). There was or indigestion; association indicated but causality
no significant difference in age between drinkers unclear: red sore eyes, impotence or loss of sex-
and non-drinkers or in usage rates between ual drive, self-reported poor health, raised cho-
patient groups or staff. The study revealed that lesterol, and loss of time and money, low
the consumption of fresh kava on a regular basis motivation and “slow/lazy” days following use,
was very common in Vanuatu. Studies by Cairney reduced alcohol consumption and related vio-
et al. (2003) on a group of current, ex and non- lence; association hypothesised: fits or seizures,
kava users among an indigenous population in melioidosis, ischaemic heart disease, protective
northern Australia found no impairment in cogni- effects for cancer; no association indicated: cog-
tive or saccade function in individuals who were nitive performance; no association suggested:
currently heavy kava users (and had been for up cognitive impairment, liver toxicity or permanent
to 18 years), nor was there any impairment in liver damage and other pneumonia; no associa-
individuals who had been heavy kava users in the tion hypothesised: hallucinations. The study of
past but had abstained for longer than 6 months. Vignier et al. (2011) found that kava drinking
Current and ex-kava users showed a higher rate was associated with suicidal behaviour among
of kava dermopathy, lower body mass index, low- young Kanaks using kava in New Caledonia but
ered blood lymphocytes and, in addition, current information on the effects of kava on mental
kava users showed elevated liver enzymes. They health in young people was lacking and require
found no evidence of brain dysfunction in heavy further investigations.
and long-term kava users. In 2000 dermopathy Chronic consumption (6 months) of large
characteristic of heavy use, abnormally low body quantities of an infusion of kava rhizome (5–6
mass index (BMI), low blood lymphocytes and cups daily) had been reported to cause anorexia,
abnormally high gamma-glutamyl transferase diarrhoea and visual disturbances (Siegel 1976).
(GGT) occurred more frequently with increased In a healthy volunteer, disturbances of visual
kava use in Arnhem Land Aboriginal communi- accommodation, such as enlargement of the
ties in the Northern Territory (Clough 2003). pupils and disturbances in oculomotor equilib-
These acute effects emerged at average consump- rium, were reported following the ingestion of
tion levels of 310–440 g/week of kava powder. large doses of kava (Garner and Klinger 1985).
The proportion of men drinking kava reached No changes were recorded in visual or stereoacu-
70 % and women 62 % from mid-1990, with ity or in ocular refractive error. Mathews et al.
20 % of the population spending unprecedented (1988) conducted a pilot health survey on kava
amounts of time (14 + hours/week) in activities usage involving 20 very heavy users of kava
where kava was consumed. (mean consumption 440 g/week), 15 were heavy
Kava being used for a wide spectrum of thera- users (310 g/week) and 4 were occasional users
peutic properties, including sedative, anxiolytic, (100 g/week). They found that in addition to
analgesic and neuroprotective effects, was one of causing acute intoxication, sedation and relax-
eight herbal supplements identified that could ation, a rash and weight loss in long-term users,
pose the greatest potential risks in surgical kava may also cause liver and renal dysfunction,
patients in the United States (Sullivan et al. hematological abnormalities and possibly pul-
2009). An Australia-wide ban on commercial monary hypertension. The conclusion that kava
importation of kava has been in place since mid- affected the liver was based on the markedly ele-
2007, but published literature on the impact of vated plasma levels of γ-glutamyl transferase
the ban had been lacking to date on the impact of observed in kava users. Heavy consumption of
the ban (Rychetnik and Madronio 2011). In a kava has been reported to lead to adverse health
182 Piperaceae

effects in some Australian Aborigines including 70-year-old man and a 52-year-old woman, with
the occurrence of pellagroid dermopathy (Cawte sebotropic drug eruption in sebaceous gland
1986, 1988). Bilia et al. (2002) reviewed nine areas induced by 3 weeks of systemic kava kava
clinical trials and found that three of these trials anti-depressant therapy. Huynh et al. (2014)
reported no adverse events, while the other six reported a 55-year-old man presented with an
studies reported gastrointestinal symptoms, tired- eruption in a sebotropic distribution after con-
ness, restlessness, tremor and headache. The suming kava kava for 3 weeks, which resolved
number of patients reporting the complaints was after discontinuation of the supplement. Shimoda
similar in the placebo groups. Aside from liver et al. (2012) found that mast cells exposed to
abnormalities or toxicity, adverse effects attrib- standardised kava aqueous extracts displayed
uted to kava extracts included gastrointestinal robust intracellular free calcium responses and
complaints, restlessness, mydriasis, allergic skin concomitant release of proinflammatory media-
reactions, dermatomyositis (Ernst et al. 2001), tors. In contrast, mast cells were refractory to
visual accommodation disorders, pupil dilation single or combinatorial stimulation with kavalac-
and disorders of oculomotor equilibrium (Singh tones, including methysticin, dihydromethysticin
and Blumenthal 1997). Toxic doses (several and kavain. The study suggested that mast cell
times the therapeutic dose of approximately activation may be a mechanistic component of
70 mg of kavalactones three times daily) could kava-related skin inflammations.
cause progressive ataxia, muscle weakness and Studies by Xia et al. (2012) suggested that
ascending paralysis (Spillane et al. 1997). kava was photocytotoxic and photogenotoxic,
Heavy kava drinkers were reported to acquire both mediated by free radicals generated during
a reversible peculiar, scaly, ichthyosiform erup- photoirradiation. They found that UVA irradia-
tion on the skin and kava dermopathy (Norton tion of kava in the presence of a lipid, methyl
and Ruze 1994). Traditional kava use at frequent linoleate, generated lipid peroxidation which was
high doses had been found to cause a reversible mediated by singlet oxygen generated during
dermatological condition known as ichthyosi- photoirradiation. Of the six major kavalactones
form ichthyosiform kava dermopathy (Süss and (yangonin; 7,8-dihydrokawain; kawain,
Lehmann 1996; Dentali 1997), known as kani- 7,8-dihydromethysticin; methysticin; and
kani in Fijian (Gounder 2006). Kava dermopathy 5,6-dehydrokawain), only 5,6-dehydrokawain
was believed to be related to interference with and yangonin-induced a low level of lipid peroxi-
cholesterol metabolism (Ruze 1990; Norton and dation. UVA irradiation of kava in human HaCaT
Ruze 1994; Dentali 1997). Ruze (1990) found skin keratinocytes induced cytotoxicity which
that the pellagroid dermopathy was not related to was mediated by oxidative stress, led to DNA
niacin deficiency and was more characteristic of strand cleavage, and produced 8-hydroxy-2′-
an acquired ichthyosis. In Fiji in 2012, over 1000 deoxyguanosine (8-OHdG) adduct. Study by the
kava participants underwent full skin examina- electron spin resonance (ESR) method revealed
tion, and kava dermopathy was a common cuta- that UVA irradiation of kava produced singlet
neous finding (Hannam et al. 2014). The clinical oxygen and carbon-centered radicals.
manifestations of kava dermopathy shared simi- Schelosky et al. (1995) reported four cases of
larities with the spectrum of autosomal recessive patients who developed clinical signs suggestive
congenital ichthyoses, predominantly lamellar of central dopamine antagonisms after exposure
ichthyosis. The pathogenesis of Kava dermopa- to various kava preparations for anxiety. These
thy may be associated with a functional defect in case histories suggested that the sedative effects
one or more cytochrome P450 enzymes impli- of kava might result from dopamine antagonistic
cated in epidermal integrity, thus mimicking the properties of kava extracts. They highlighted the
genetic defect as seen in lamellar ichthyosis type pyramidal side effects of kava preparations and
3. Jappe et al. (1998) described two patients, a cautioned their use particularly in elderly patients.
Piper methysticum 183

Meseguer et al. (2002) reported a 45-year-old In Tubuai, a maceration of young kava shoot is
female with severe parkinsonism induced by taken orally for inflammation of the genito-
kava kava. The patient, who had a family history urinary system (Aitken 1930) In the Pacific
of essential tremor, developed severe and persis- Islands, kava is taken as a drink for gonorrhoea
tent parkinsonism after days of treatment with and chronic cystitis (Steinmetz 1960). In Papua
kava extract for anxiety. The symptoms improved New Guinea, kava is used as an anaesthetic and
with anticholinergics. They asserted that kava stimulant for lactation (Steinmetz 1960); the bark
derivatives could produce severe parkinsonism in of scraped and masticated root is used to relive
individuals with genetic susceptibility. sore throats, juice from leaves is used to treat cuts
A case of rhabdomyolysis temporally related and drunk as tonic (Holdsworth 1977). In certain
to the ingestion of a large amount of kava was tribes of Papua New Guinea, women drink fresh
reported by Bodkin et al. (2012) in addition to an masticated kava as an anaesthetic, when they are
earlier case of rhabdomyolysis related to the being tattooed (Steinmetz 1960). They also drink
ingestion of kava. Their patient developed peak large quantities when they are pregnant, espe-
creatine phosphokinase levels in excess of 30 000 cially just before delivery, to stimulate milk pro-
U/L but had no significant renal damage. In May duction. In Irian Jaya, the internal skin of the
2013, a case of hepatitis A infection was reported kava plant is used for toothache (Aufenanger and
to a Western Australian regional public health Hötker 1940). In Tahiti, according to Sterly
unit, with infection acquired in Fiji (Parker et al. (1970), the masticated root is considered a rem-
2014). Following this, two further cases were edy for gonorrhoea. In American Samoa, kava
linked to the index case by kava drinking and one roots together with fruit of polo fe’u (chilli,
further case was a household contact of a second- Capsicum annuum), leaves of fisoa (soap bush,
ary case. This outbreak highlighted that the prep- Colubrina asiatica) and inner bark of moli’aina
aration of kava drink and/or the use of a common (orange, Citrus sinensis) are taken internally for
drinking vessel could be a vehicle for the trans- gonorrhoea; kava leaves are taken internally for
mission of hepatitis A virus. mumu tuala uli (swollen head, sore eyes, cold
sweat, dizziness, numbness of legs), and the inner
bark with juice of fasa (screw-pine, Pandanus
Traditional Medicinal Uses tectorius) is taken internally for tulita fasia (uri-
nary tract infection) Harrington and Scotese
Kava has been used in social and ceremonial life (2001).
in the Pacific islands from ancient times for the In Hawaii, awa root is used to treat difficulties
soporific and narcotic effects (Bilia et al. 2002). in urination (Handy et al. 1934; Titcomb 1948)
In Polynesia, about thirty syndromes were treated and irritation of the genito-urinary system (Handy
with kava-based preparations (Zepernick 1972). et al. 1934); masticated awa and drinking awa are
The traditional medicinal uses of kava include employed for feminine puberty syndromes and
treatment of gonorrhoea, syphilis, and cystitis; weakness, painful migraine, headache, macera-
induction of muscle relation and sleep (Ernst tion of awa for vaginal prolapus, maceration of
et al. 2001); and treatment of boils, asthma, head- masticated awa diluted, boiled and taken orally
ache and urinary infections (Hirsch 2000). In the for general weakness (Titcomb 1948); awa-based
Pacific Oceania, kava is held in high esteem pri- medication is used for menstrual problems and
marily as a ritual offering or a ritualised form of dysmenorrhoea, and masticated awa is drank to
payment (Lebot and Cabalion 1988). To treat prevent risk of infection (Handy et al. 1934);
injury caused by fish spines, dry kava root is maceration of masticated awa is diluted, boiled
burnt and the injury exposed to the smoke and taken orally for chills and sleeping problems
generated. (Titcomb 1948: Handy et al. 1934; Handy 1940;
184 Piperaceae

Hänsel 1968); masticated awa or as a drink is sumed by women as a fortifying drink, laxative
employed for headaches (Handy et al. 1934; and diuretic. In pregnancy, small amount taken is
Handy 1940; Titcomb 1948); maceration of rhi- said to facilitate delivery. During breast feeding,
zome is taken orally for rheumatism and against awa is taken to stimulate milk production. Kava
fat intake (Handy 1940; Titcomb 1948); awa is used medicinally also for kidney and bladder
preparation is employed for the irritation of respi- troubles and as a strong sudorific. Though like
ratory tract, a maceration of rhizome is taken all other drugs, if taken in excess it has bad
orally for gastro-intestinal upsets, masticated awa results on account of its excessive action on the
is drank for pulmonary pains, awa is used for cer- skin, and may even super-induce elephantiasis,
tain skin disease and masticated rhizome as poul- that terrible complaint, so prevalent in Fijian vil-
tice against suppuration (Titcomb 1948). Awa lages. In Fiji, kava leaves are softened in fire and
drink is administered to children to calm their applied as poultice against suppurations
nervousness (Titcomb 1948), juice from rhizome (Degener 1949). Fijian kava is used in traditional
is used in medication for tuberculosis, medica- medicine against urinary tract infections and
tion containing masticated rhizome used exter- asthma (Folmer et al. 2006). Japanese newspaper
nally for leprosy (Degener 1945) and masticated in early 1985 reported virtues of Fijian yaqona
rhizome is used in poultice for skin disease as a remedy against colds and coughing (Lebot
(Hänsel 1968). Awa leaves are placed in-situ to and Cabalion 1988)
provoke abortion (Handy et al. 1934), and fumi- Ingestion of kava is also reported to aid in first
gation with leaves is used for chill and general stage of diarrhoea (Thomson 1908; Sterly 1970).
disease treatment (Handy 1940). In Motalava, Vanuatu, kava roots are used in
In Hawaii, Awa is taken as a therapy for sooth- drink to cure constipation and used for conjuncti-
ing the nerves, relaxing fatigue-stiff muscles, vitis (Vienne 1981). Eyes are washed with juice
general debility and inducing sleep and a treat- squeezed from kava leaves.
ment for excessive fat for restoring the body to
normal fitness (Titcomb 1948). Awa treatment
was also said to overcome scaly and ulcerous Other Uses
skin. Mouritz (1916) stated that an alcoholic awa
solution injected into the skin causes anaesthesia, The use of kava over the centuries is in the form
followed by paralysis of the peripheral nerves for of a traditional ethnic beverage forming an inte-
several hours. He added that both Hawaiian and gral part of social gatherings and ceremonies to
haole (foreign) doctors use it for venereal and increase kinship and amiability, especially in the
kidney diseases, and, in alcoholic solution or as Polynesian and Melanesian societies, and is
an unguent, in affections of the skin, including widely consumed on a daily basis.
leprosy. Handy et al. (1934) stated that bits of Kava leaf/root/stem extract is used as cos-
awa root chewed at frequent intervals and awa metic skin-conditioning agents at concentrations
leaves wrapped around the head were said to pro- from 0.0001 to 0.1 % (Robinson et al. 2009).
tect against contagious disease and to cure head- Data is still wanting on whether kava-derived
ache. Awa leaves, stuffed into the vagina, were ingredients would be substantially absorbed
said to induce miscarriage. Jarrett (1933) men- through the skin.
tioned that awa was extensively used in Germany Kava had been reported to have antifungal and
prior to the World War, in the manufacture of cer- herbicidal property.
tain drugs and medicines. Kava showed a strong inhibition on growth of
In Fiji, yaqona is considered a powerful dia- barnyardgrass (Echinochloa crus-galli), mono-
phoretic (Parham 1939, 1941). It is also con- choria (Monochoria vaginalis) and knotgrass
Piper methysticum 185

(Paspalum distichum), some of the most harmful Abu N, Akhtar MN, Yeap SK, Lim KL, Ho WY, Zulfadli
AJ, Omar AR, Sulaiman MR, Abdullah MP, Alitheen
paddy weeds (Xuan et al. 2003). Kava completely
NB (2014) Flavokawain A induces apoptosis in
controlled emergence of monochoria and barn- MCF-7 and MDA-MB231 and inhibits the metastatic
yardgrass at a treated dose of 0.5 and 1.0 g, process in vitro. PLoS One 9(10):e105244
respectively. Application of kava at 1 tonne/ha 6 Achenbach H, Karl W (1970) Uber die isolierung von
zwei neuen pyrrolidinen aus rauschpfeffer. Chem Ber
days after saturating paddy soil with water was
103:2535–2540. (In German)
an effective treatment. This caused around 80 % Achenbach H, Karl W (1971) Untersuchung der sauren
reduction of natural paddy weed growth and des rauschpfeffers (Piper methysticum Forst.). Chem
increased tillering and root number of rice. In Ber 103:2535–2540
Achenbach H, Wittman G (1970) Dihydrokawain-5-ol,
addition, kava significantly inhibited growth of
ein neuer alkohol aus rauschpfeffer (Piper methysti-
the five fungi: Fusarium solani, Pyricularia gri- cum Forst.). Tetrahedron Lett 11(37):3259–3262
sea, Rhizopus stolonifer, Taphrina deformans Achenbach H, Karl W, Smith S (1971) Zur gaschro-
and Thanatephorus cucumeris. The effect on R. matograpischen trennung der kawa-lactone-(+)-
5,6,7,8-tetrahydro-yangonin, ein neus kawa-lacton aus
stolonifer was the greatest and T. cucumeris and
rauschpfeffers. Chem Ber 104:2688–2693
P. grisea were the second most affected. Aqueous Achenbach H, Karl W, Regel W (1972) 11-Hydroxy-12-
extract of kava roots showed high allelopathic methoxy-dihydrokawain und 11,12-dimethoxy-
potential and strongly suppressed germination dihydrokawain, zwei neue kawa-lactone aus
rauschpfeffer (Piper methysticum Forst). Chem Ber
and growth of lettuce, radish, barnyardgrass
and Monochoria (Xuan et al. 2006). The six Aitken RT (1930) Ethnology of Tubaui. Bernice P Bishop
major lactones desmethoxyyagonin, kavain, Mus Bull 70:42
7,8-dihydrokavain, yagonin, methysticin and Almeida JC, Grimsley EW (1996) Coma from the health
food store: interaction between kava and alprazolam.
dihydromethystici in kava roots showed great
Ann Int Med 125(11):940–941
herbicidal and antifungal activities. Growth of Anke J, Ramzan I (2004) Pharmacokinetic and pharma-
lettuce and barnyardgrass were significantly codynamic drug interactions with Kava (Piper
inhibited at 1–10 ppm, and four plant fungi methysticum Forst. f.). J Ethnopharmacol 93(2–3):
including Colletotrichum gloeosporioides,
Aufenanger H, Hötker G (1940) Die Gende in
Fusarium solani, Fusarium oxysporum and Zentralneuguinea, vom leben und denken eines Papua-
Trichoderma viride were significantly inhibited Stammes im Bismarck gebirge. [The gende of central
at 10–50 ppm (Xuan et al. 2006). New Guinea, life and thought of a Papuan branch in
the Bismarck mountains.] Erganzungsbande zur
Ethnographie Neuguineas 1. Missionsdruckerei St.
Gabriel, Wien-Modling. 209 pp
Comments Backhauss C, Krieglstein J (1992) Extract of kava (Piper
methysticum) and its methysticin constituents protect
brain tissue against ischemic damage in rodents. Eur
Kava is readily propagated from mature, freshly
J Pharmacol 215(2–3):265–269
harvested entire stems or 1–4 node sections of Barrau J (1957) A propos du Piper methysticum Forster.
stems, or by division of the root-mass, or by J Agric Trop Bot Appl 4:270–273. (In French)
removal of offsets from the root ball of the Baum SS, Hill R, Rommelspacher H (1998) Effect of kava
extract and individual kavapyrones on neurotransmit-
ter levels in the nucleus accumbens of rats. Prog
Neuropsychopharmacol Biol Psychiatry
Selected References Behl M, Nyska A, Chhabra RS, Travlos GS, Fomby LM,
Sparrow BR, Hejtmancik MR, Chan PC (2011) Liver
toxicity and carcinogenicity in F344/N rats and
Abourashed EA, Khan IA (2000) Microbial transforma-
B6C3F1 mice exposed to Kava Kava. Food Chem
tion of kawain and methysticin. Chem Pharm Bull
Toxicol 49(11):2820–2829
Bhate H, Gerster G, Gracza E (1989) Orale prämedikation
Abu N, Ho WY, Yeap SK, Akhtar MN, Abdullah MP,
mit zubereitungen aus Piper methysticum bei
Omar AR, Alitheen NB (2013) The flavokawains:
operativen eingriffen in epiduralanästhesie.
uprising medicinal chalcones. Cancer Cell Int
Erfahrungsheilkunde 6:339–345. (In German)
186 Piperaceae

Bilia AR, Gallori S, Vincieri FF (2002) Kava-kava and Borsche W, Walter C (1927) Untersuchungen über die
anxiety: growing knowledge about the efficacy and bestandteile der kawawurzel. V. Syntheses des yan-
safety. Life Sci 70(22):2581–2597 gonols. Chem Ber 60:2112. (In German)
Bobeldijk I, Boonzaaijer G, Spies-Faber EJ, Vaes WH Borsche W, Meyer CH, Peitzsch W (1927a)
(2005) Determination of kava lactones in food supple- Untersuchungen über die bestandteile der kawawur-
ments by liquid chromatography-atmospheric pres- zel. VI. Die constitution des methystizins. Chem Ber
sure chemical ionisation tandem mass spectrometry. 60:2113–2120. (In German)
J Chromatogr A 1067(1–2):107–114 Borsche W, Rosenthal W, Meyer CH (1927b)
Bodkin R, Schneider S, Rekkerth D, Spillane L, Kamali Untersuchungen über die bestandteile der kawawur-
M (2012) Rhabdomyolysis associated with kava zel. IV. Über die synthese eines kawasaure-methyesters
ingestion. Am J Emerg Med 30(4):635.e1-3 und zweier isomer des methystizins. Chem Ber
Boonen G, Beck MA, Häberlein H (1997) Contribution to 60:1135–1139. (In German)
the quantitative and enantioselective determination of Brown AC, Onopa J, Holck P, Kaufusi P, Kabasawa D,
kavapyrones by high-performance liquid chromatog- Craig WJ, Dragull K, Levine AM, Baker JD (2007)
raphy on ChiraSpher NT material. J Chromatogr B Traditional kava beverage consumption and liver func-
Biomed Sci Appl 702(1):240–244 tion tests in a predominantly Tongan population in
Boonen G, Ferger B, Kuschinsky K, Häberlein H (1998) Hawaii. Clin Toxicol (Phila) 45(5):549–556
In vivo effects of the kavapyrones (+)-dihydrome- Brüggemann F, Meyer HJ (1962) Die analgetische
thysticin and (+/−)-kavain on dopamine, 3,4- wirkung der kawa-inhaltsstoffe dihydrokawain und
dihydroxyphenylacetic acid, serotonin and 5-hydroxy- dihydromethysticin. Arzneim Forsch 12:407–409. (In
indoleacetic acid levels in striatal and cortical brain German)
regions. Planta Med 64(6):507–510 Bruner NR, Anderson KG (2009) Discriminative-stimulus
Borsch W, Blount BK (1930) Untersuchungen über die and time-course effects of kava-kava (Piper methysti-
bestandteile der kawawurzel. XI. Synthese der methys- cum) in rats. Pharmacol Biochem Behav
tizinsaure und der kawasäure. Chem Ber 63:2418– 92(2):297–303
2420. (In German) Buckley JP, Furgiuele AR, O’Hara MJ (1967)
Borsch W, Blount BK (1932) Untersuchungen über die Pharmacology of kava. In: Efron DH (ed)
bestandteile der kawawurzel. XII. Über yangonlakton Ethnopharmacologic search for psychoactive drugs,
und triacetäure. Chem Ber 65:820–828. (In German) United States Public Health Service publication no.
Borsch W, Blount BK (1933) Untersuchungen über die 1645. United States Public Health Service,
bestandteile der kawawurzel. XIII. Über einige neue Washington, DC
stoffe aus technischen kawaharz. Chem Ber 66:803– Cagnacci A, Arangino S, REnzi A, Zanni AL, Malmusi S,
806. (In German) Volpe A (2003) Kava- kava administration reduces
Borsch W, Lewinsohn M (1933) Untersuchungen über die anxiety in perimenopausal women. Maturitas
bestandteile der kawawurzel. XIV. Über cinnamol- 44(2):103–109
essigester. Chem Ber 66:1792–1801. (In German) Cairney S, Maruff P, Clough AR (2002) The neurobehav-
Borsche W (1927) Untersuchungen über die bestandteile ioural effects of kava. Aust N Z J Psychiatry
der kawawurzel. III. Über die katalytische hydrierung 36(5):657–662
des methystizins. Chem Ber 60:982–984. (In German) Cairney S, Maruff P, Clough AR, Collie A, Currie J,
Borsche W, Bodenstein CK (1929) Untersuchungen uber Currie BJ (2003) Saccade and cognitive impairment
die bestandteile der kawawurzel. IX. Die syntheses associated with kava intoxication. Hum
des yangonins. Chem Ber 62:2515–2523. (In German) Psychopharmacol 18(7):525–533
Borsche W, Gerhard M (1914) Untersuchungen über die Campo JV, McNabb J, Perel JM, Mazariegos GV,
bestandteile der kawawurzel.I. Über yangonin. Chem Hasegawa SL, Reyes J (2002) Kava-induced fulmi-
Ber 47:2902–2918. (In German) nant hepatic failure. J Am Acad Child Adolesc
Borsche W, Peitzsch W (1929a) Untersuchungen über die Psychiatry 41(6):631–632
bestandteile der kawawurzel. VII. Über kawasäure. Cawte J (1986) Parameters of kava used as a challenge to
Chem Ber 62:368–373. (In German) alcohol. Aust NZ J Psychiatry 20(1):70–76
Borsche W, Peitzsch W (1929b) Untersuchungen über die Cawte J (1988) Macabre effects of a cult for kava. Med
bestandteile der kawawurzel. VII. Über pseudo- J Aust 148(11):545–546
methystizin. Chem Ber 62:360–367. (In German) Cheng D, Lidgard RO, Duffield PH, Duffield AM (1988)
Borsche W, Peitzsch W (1930) Untersuchungen über die Identification by methane chemical ionization gas
bestandteile der kawawurzel. X. Über kawain and chromatography/mass spectrometry of the products
dihydrokawain. Chem Ber 63:2414–2417. (In obtained by steam distillation and aqueous acid extrac-
German) tion of commercial Piper methysticum. Biomed
Borsche W, Roth A (1921) Untersuchungen über die bes- Environ Mass Spectrom 17(5):371–376
tandteile der kawawurzel.II. Über das kawaharz. Chienthavorn O, Smith RM, Wilson ID, Wright B, Lenz
Chem Ber 54:2229–2235. (In German) EM (2005) Superheated water chromatography-
Piper methysticum 187

nuclear magnetic resonance spectroscopy of kava lac- Duffield PH, Jamieson DD (1991) Development of toler-
tones. Phytochem Anal 16(3):217–221 ance to kava in mice. Clin Exp Pharmacol Physiol
Clouatre DL (2004) Kava kava: examining new reports of 18(8):571–578
toxicity. Toxicol Lett 150(1):85–96 Duffield AM, Lidgard RO (1986) Analysis of kava resin
Clough AR (2003) Enough! or too much. What is ‘exces- by gas chromatography and electron impact and meth-
sive’ kava use in Arnhem Land? Drug Alcohol Rev ane negative ion chemical ionization mass spectrome-
22(1):43–51 try. New trace constituents of kava resin. Biol Mass
Clough AR, Bailie RS, Currie B (2003a) Liver function Spectrom 13:621–626
test abnormalities in users of aqueous kava extracts. Duffield AM, Lidgard RO, Low GKC (1986) Analysis of
J Toxicol Clin Toxicol 41(6):821–829 the constituents of Piper methysticum by gas chroma-
Clough AR, Jacups SP, Wang Z, Burns CB, Bailie RS, tography methane chemical ionization mass spectrom-
Cairnery SJ, Collie A, Guyula T, McDoanld SP, Currie etry. New constituents of kava resin. Biol Mass
BJ (2003b) Health effects of kava use in an eastern Spectrom 13:305–313
Arnhem Land aboriginal community. Intern Med Duffield AM, Jamieson DD, Lidgard RO, Duffield PH,
J 33(8):336–340 Bourne DJ (1989a) Identification of some human uri-
Côté CS, Kor C, Cohen J, Auclair K (2004) Composition nary metabolites of the intoxicating beverage kava.
and biological activity of traditional and commercial J Chromatogr A 475(2):273–281
kava extracts. Biochem Biophys Res Commun Duffield PH, Jamieson DD, Duffield AM (1989b) Effect
322(1):147–152 of aqueous and lipid-soluble extracts of kava on the
Cuzent G (1981) Rapport sur la composition chimique de conditioned avoidance response in rats. Arch Int
la kavahine. CR Acad Sci Paris 52(5):205–206. (In Pharmacodyn Ther 301:81–90
French) Dutta CP, Ray LPK (1973) Further studies on the structure
Davies LP, Drew CA, Duffield P, Johnston GA, Jamieson of flavokawain C. A new chalcone isolated from Piper
DD (1992) Kava pyrones and resin: studies on methysticum. Indian Sci Congress Assoc Proc
GABAA, GABAB and benzodiazepine binding sites 60:121–122
in rodent brain. Pharmacol Toxicol 71(2):120–126 Dutta CP, Ray LPK, Chatterjee A, Roy DN (1972)
de Jager LS, Perfetti GA, Diachenko GW (2004) LC-UV Constituents of Piper methysticum. Phytochemistry
and LC-MS analysis of food and drink products con- 11(9):2891–2892
taining kava. Food Addit Contam 21(10):921–934 Dutta CP, Ray LPK, Chatterjee A, Roy DN (1973)
De Leo V, La Marca A, Lanzetta D, Palazzi S, Torricelli Constituents of flavokawain C. Piper methysticum. Ind
M, Facchini C, Morgante G (2000) Assessment of the J Chem 11(5):509–510
association of Kava-Kava extract and hormone Dutta CP, Ray LPK, Chatterjee A, Roy DN (1976) Studies
replacement therapy in the treatment of postmeno- on the genus Piper. V. Chemical investigation of Piper
pause anxiety. Minerva Ginecol 52(6):263–267. (In methysticum Forst. (Piperqceae). Structure and syn-
Italian) thesis of flavokawain C. J Ind Chem Soc
Degener O (1945) Plants of Hawaii National Park, 2nd 53(12):1194–1197
edn. Edwards Brothers, Waialua, Oahu, Hawaii Dutta CP, Som UK (1978) Studies on the genus Piper –
Degener O (1949) Naturalists’ South Pacific expedition: VIII. Further studies on the roots of Piper methysticum
Fiji. Paradise of the Pacific, Ltd, Honolulu. 303 pp Forst. J Indian Chem Soc 55:932–934
Denham A, McIntyre M, Whitehouse J (2002) Kava – the Duve RN (1976) Highlight of the chemistry and pharma-
unfolding story: report on a work-in-progress. J Altern cology of yaqona, Piper methysticum. Fiji Agric
Complement Med 8(3):237–263 J 38(2):81–84
Dentali SJ (1997) Herb Safety Review, Piper methysticum Duve RN (1981) Gas-liquid chromatographic determina-
Forster f. (Piperaccae). Herb Research Foundation, tion of major constituents of Piper methysticum.
Boulder Analyst 106:160–165
Dharmaratne HRW, Nanayakkara NPD, Khan IA (2002) Duve RN, Prasad J (1983) Changes in the chemical com-
Kavalactones from Piper methysticum and their 13C position of yaqona Piper methysticum with time. Fiji
NMR spectroscopic analyses. Phytochemistry Agric J 45(2):45–50
59:429–433 Emser W, Bartylla K (1991) Verbesserung der schlafqual-
Dinh LD, Simmen U, Bueter KB, Bueter B, Lundstrom K, ität. Zur wirkung von kava-extrakt WS 1490 auf das
Schaffner W (2001) Interaction of various Piper schlafmuster bei gesunden. TW Neurol Psychiatr
methysticum cultivars with CNS receptors in vitro. 5:636–642. (In German)
Planta Med 67(4):306–311 Ernst E, Pittler MH, Stevinson C, White AR, Eisenberg D
DiSilvestro RA, Zhang W, DiSilvestro DJ (2007) Kava (2001) The desktop guide to complementary and alter-
feeding in rats does not cause liver injury nor enhance native medicine. Mosby, Edinburgh
galactosamine-induced hepatitis. Food Chem Toxicol Eskander RN, Randall LM, Sakai T, Guo Y, Hoang B, Zi
45(7):1293–1300 X (2012) Flavokawain B, a novel, naturally occurring
Dragull K, Yoshida WY, Tang CS (2003) Piperidine alka- chalcone, exhibits robust apoptotic effects and induces
loids from Piper methysticum. Phytochemistry G2/M arrest of a uterine leiomyosarcoma cell line.
63(2):193–198 J Obstet Gynaecol Res 38(8):1086–1094
188 Piperaceae

Fackelmann KA (1992) The history, chemistry and botany Gleitz J, Beile A, Wilkins P, Ameri A, Peters T (1997)
of the mind-altering kava plant. Sci News Antithrombotic action of the kava pyrone (+)-kavain
141:424–425 prepared from Piper methysticum on human platelets.
Feltenstein MW, Lambdin LC, Ganzera M, Ranjith H, Planta Med 63(1):27–30
Dharmaratne W, Nanayakkara NP, Khan IA, Sufka KJ Gounder R (2006) Kava consumption and its health
(2003) Anxiolytic properties of Piper methysticum effects. Pacific Health Dialog 13(2):131–135
extract samples and fractions in the chick social- Gow PJ, Connelly NJ, Hill RL, Crowley P, Angus PW
separation-stress procedure. Phytother Res (2003) Fatal fulminant hepatic failure induced by a
17(3):210–216 natural therapy containing kava. Med J Aust
Folmer F, Blasius R, Morceau F, Tabudravu J, Dicato M, 178(9):442–443
Jaspars M, Diederich M (2006) Inhibition of Grace RF (2003) Kava drinking in Vanuatu – a hospital
TNFalpha-induced activation of nuclear factor kappaB based survey. Pacific Health Dialog 10(2):41–44
by kava (Piper methysticum) derivatives. Biochem Gracza L, Ruff P (1986) Uber die hoheren alkohole von
Pharmacol 71(8):1206–1218 Piperis methystici rhizome: aliphatic and alicyclic
Frater AS (1958) Medical aspects of kava. Trans Proc Fiji alcohols of Piperis methystici rhizome. Arch Pharm
Soc 5(2):31–39 319:475–477
Fu S, Rowe A, Ramzan I (2012) Kavalactone metabolism Grunze H, Langosch J, Schirrmacher K, Bingmann D,
in the isolated perfused rat liver. Phytother Res Von Wegerer J, Walden J (2001) Kava pyrones exert
26(12):1813–1816 effects on neuronal transmission and transmembrane-
Furgiuele AR, Kinnard WJ, Aceto MD, Buckley JP ous cation currents similar to established mood stabi-
(1965) Central activity of aqueous extracts of Piper lizers – a review. Prog Neuropsychopharmacol Biol
methysticum (kava). J Pharm Sci 54(2):247–252 Psychiatry 25(8):1555–1570
Garner LF, Klinger JD (1985) Some visual effects caused Guérin J-C, Réveillère HP (1984) Activité antifongique
by the beverage kava. J Ethnopharmacol 13(3): d’extraits végétaux à usage thérapeutique. I. Étude de
307–311 41 extraits sur 9 souches fongiques. Ann Pharm Fr
Garrett KM, Basmadjian G, Khan IA, Schaneberg BT, 42:553–559. (In French)
Seale TW (2003) Extracts of kava (Piper methysti- Guo L, Li Q, Xia Q, Dial S, Chan PC, Fu P (2009)
cum) induce acute anxiolytic-like behavioral changes Analysis of gene expression changes of drug metabo-
in mice. Psychopharmacology (Berl) 170(1):33–41 lizing enzymes in the livers of F344 rats following oral
Gastpar M, Klimm HD (2003) Treatment of anxiety, ten- treatment with kava extract. Food Chem Toxicol
sion and restlessness states with Kava special extract 47(2):433–442
WS 1490 in general practice: a randomized placebo- Guo L, Shi Q, Dial S, Xia QS, Mei N, Li QZ, Chan PC, Fu P
controlled double-blind multicenter trial. (2010) Gene expression profiling in male B6C3F1 mouse
Phytomedicine 10(8):631–639 livers exposed to kava identifies – changes in drug metab-
Gautz LD, Kaufusi P, Jackson MC, Bittenbender HC, olizing genes and potential mechanisms linked to kava
Tang CS (2006) Determination of kavalactones in toxicity. Food Chem Toxicol 48(2):686–696
dried kava (Piper methysticum) powder using near- Häberlein H, Boonen G, Beck MA (1997) Piper methysti-
infrared reflectance spectroscopy and partial least- cum: enantiomeric separation of kavapyrones by high
squares regression. J Agric Food Chem performance liquid chromatography. Planta Med
54(17):6147–6152 63(1):63–65
Gessner B, Cnota P (1994) Untersuchung der vigilanz Halzl JS, Juretzek W, StahlBiskup E (1993) Piper. In: von
nach applikation von kava-kava-extrakt, diazepam Bruchhausen F, et al (12 others) (eds) Hagers
oder placebo. (Extract of the kava-kava rhizome in Handbuch der Pharmazeutischen Praxis, 5th edn.
comparison with diazepam and placebo). Z Phytother Springer, Berlin. 5:52–59
15:30–37. (In German) Handy ESC (1940) The Hawaian planter. Volume I. His
Gleitz J, Beile A, Peters T (1995) (+/−)-Kavain inhibits plants, methods and areas of cultivation, Bernice
veratridine-activated voltage-dependent Na(+)- P. Bishop Mus Bull 161. 227pp
channels in synaptosomes prepared from rat cerebral Handy ESC, Pukui MK, Livermore K (1934) Outline of
cortex. Neuropharmacology 34(9):1133–1138 Hawaiian physical therapeutics, Bernice P. Bishop
Gleitz J, Beile A, Peters T (1996a) (+/−)-kavain inhibits Museum Bull 126. 51 pp
the veratridine- and KCl-induced increase in intracel- Hannam S, Murray M, Romani L, Tuicakau M, Whitfeld
lular Ca2+ and glutamate-release of rat cerebrocortical MJ (2014) Kava dermopathy in Fiji: an acquired ich-
synaptosomes. Neuropharmacology 35(2):179–186 thyosis? Int J Dermatol 53(12):1490–1494
Gleitz J, Friese J, Beile A, Ameri A, Peters T (1996b) Hänsel R (1968) Characterisation and physiological activ-
Anticonvulsive action of (+/−)-kavain estimated from ity of some kava constituents. Pacific Sci 22:293–313
its properties on stimulated synaptosomes and Hänsel R, Klaproth L (1966) Isolierung von 11-methoxy-
Na + channel receptor sites. Eur J Pharmacol 315(1): yangonin aus der kawawurzel. Arch Pharm 299:
89–97 503–506
Piper methysticum 189

Hänsel R, Schulz J (1973) Synthese von racem 5,6-cis- Holm E, Staedt U, Heep J, Koetsik C, Behne F, Kaske S,
und trans-kawain-5-ol. Chem Ber 106:570–575 Mennicke I (1991) Untersuchungen zum wirkung-
Hänsel R, Baehr P, Elich J (1961) Isolierung und charak- sprofil von d,l-kavain. Zerebrale angriffsorte und
terisierung von zwei bisher unbekannten farbstoffen schlaf-wach-rhythmus im tierexperiment.(The action
des kawa-rhizoms eine vorläufige m