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435© Springer Nature Switzerland AG 2020
J. Novak, W.-D. Blüthner (eds.), Medicinal, Aromatic and Stimulant Plants,
Handbook of Plant Breeding 12, https://doi.org/10.1007/978-3-030-38792-1_13
Chapter 13
Petroselinum crispum (Mill.) Nyman
(Parsley)
FrankMarthe
13.1 Botany (Taxonomy, Origin, Distribution, Cytology,
Plant Description)
Parsley (Petroselinum crispum) belongs to the family Apiaceae (syn. Umbelliferae).
Many species of this family are used for long time as medicinal and aromatic plants
as well as for vegetables.
According to Hanelt (2001) the genus Petroselinum J.Hill, Brit. herb. (1756)
424, includes two species, P. crispum and Petroselinum segetum [L.] W.D.J.Koch
(corn parsley). P. crispum occurs naturally in Western Europe from the British Isles
to the Iberian Peninsula. Parsley comprises considerable variability according to its
long and different use. Lené described it as Apium petroselinum. Actually, parsley
belongs to genus Petroselinum with the name Petroselinum crispum (Mill.) Nyman,
Consp. . eur. 2 [1879] 309 (nomen in syn.) ex A.W.Hill, Hand-list herb. pl. Kew
ed. 3 (1925) 122 Airy-Shaw in Kew Bull. (1938) 256).
The name of the genus Petroselinum was derived from the Greek word petra
(Πέτϱα) for rock or stone and Latin word selīnum for plants growing on stony soil.
The name of the species crispum comes from the Latin crispus and means curled.
Common names for parsley are Arabian, سنودقب (madanous); Chinese, 芫荽 (yánsuī);
French, persil, turnip-rooted parsley (persil à grosse racine); German, Petersilie,
turnip-rooted parsley (Wurzelpetersilie); Italian, prezzemolo; Persian, یرفعج (jaaf-
eri); Portuguese, salsa; Russian, петрушка (petruška); Spanish, perejil; Turkish,
maydanoz; Vietnamese, Mùi tây.
Common names combined with parsley exist meaning other species than
P. crispum (e.g. Chinese or Japanese parsley for coriander (Coriandrum sativum
L.)). Sometimes mitsuba or Japanese parsley (Cryptotaenia canadensis [L.] DC. ssp.
F. Marthe (*)
Julius Kuehn Institute, Federal Research Centre for Cultivated Plants, Quedlinburg, Germany
e-mail: frank.marthe@julius-kuehn.de
436
japonica [Hassk.] Hand.-Mazz.) from Japan is confused with parsley. Beaked
parsley, French parsley or gourmet parsley refers to chervil (Anthriscus cerefolium
[L.] Hoffm.). Vietnamese parsley sometimes means Vietnamese coriander or rau
răm (Polygonum odoratum Lour.) or water dropwort or cần nước (Oenanthe javan-
ica [Blume] DC.) (Small 1997).
Two convarieties are differentiated in parsley. The convar. radicosum (Alef.)
Danert in Mansf. Verzeichnis (1959) 322 with eatable swollen taproots and convar.
crispum where the roots are not eaten. In all populations of convar. radicosum leaves
are not crispy; they belong to var. tuberosum (Bernh.) Crov. Non-crispy accessions
of var. erfurtense Danert were available from gene banks. Parsley roots of convar.
radicosum are used as vegetable. Among convar. crispum different leaf types exist:
accessions with crispy leaves attributed to var. crispum, with non-crispy leaves to
var. vulgare (Nois.) Danert and var. neapolitanum Danert also known as Neapolitan
or Italian parsley with eshy and longer petioles, respectively (Fig.13.1). In the
Mediterranean region, some cultivars are grown for these edible leaf stalks (Small
1997). Accessions of convar. crispum var. silvestre (Alef.) Danert, the wild type of
parsley, were also not available from gene banks (Danert 1959).
The origin of parsley is doubtful, but comes probably from the eastern part of
Mediterranean area or Western Asia, where natural variability is highest (Hanelt and
IPK 2001). North Africa has also high variability (Ipor and Oyen 1999). The annual
accessions, which do not require vernalization, come from this region. Actually,
Fig. 13.1 Parsley (Petroselinum crispum); (a) at leaves of convar. crispum var. vulgare; (b)
crisped leaves of convar. crispum var. crispum; (c) taproots of convar. radicosum var. tuberosum
F. M ar the
437
parsley is grown at the subtropical and temperate zone around the world and escaped
from cultivation and naturalized on many places around the world. It grows also
naturally in the western part of the Mediterranean area and so it overlaps with the
region of P. segetum. There is no information about natural or articial bastards of
both species.
The chromosome number of parsley is 2n=2x=22 and of P. segetum 2n=18
(Darlington and Wylie 1955). Own evaluation was done by ow cytometry for level
of ploidy of 50 parsley accessions obtained from the federal ex situ collection of
agricultural and horticultural plants at the Leibniz Institute of Plant Genetics and
Crop Plant Research at Gatersleben, Germany (IPK). For all of them the diploid
status (2n=2x = 22) was detected. The DNA amount of diploid Petroselinum
crispum was measured as 2C=4.0pg (Bennett and Leitch 1995).
Parsley is mostly a biennial species. It can grow also semi-perennial in green-
house or warmer regions. Annual accessions also exist without requirement of ver-
nalization. Biennial accessions form in the rst vegetation period the rosette of
leaves. The length of the tripinnate leaves with numerous leaets varies in a wide
range inside the species from around 8 to 25cm. Leaves of var. crispum are shorter
and the longest are found in var. neapolitanum.
In relation to other medicinal and aromatic plants, genetic resources of parsley
are considerably represented in ex situ collections worldwide. The biggest parsley
sets exist in the following countries: Spain, 351 accessions; Germany, Federal ex
situ Gene Bank at Leibniz Institute of Plant Genetics and Crop Plant Research
(IPK), 242 accessions; United States, 249 accessions (USDA 2019); Poland, 229
accessions; Hungary, 163 accessions; Russian Federation, N.I.Vavilov All-Russian
Research Institute of Plant Industry (VIR), 66 accessions; Ukraine, 163 accessions;
Portugal, 113 accessions; Bulgaria, 82 accessions; United Kingdom, 67 accessions;
Romania, 57 accessions; and Czech Republic, 40 accessions (Eurisco 2019). In the
collections, many landraces exist from long traditions of use at geographically dif-
ferent places as well as old varieties.
13.2 Economical Use
Parsley has been used as a crop plant for more than 2500years. The rst mentioning
was made by Theophrastus (322 BC), an ancient Greek botanist and physician
(Small 1997). Hippocrates (ca 460–ca 370 BC) applied parsley as a diuretic. The
ancient Greeks held parsley sacred, using it not only to adorn victors of athletic
contests but also for decorating tombs. It was never brought to the dinner table. The
ancient romans brought it to Central Europe and in the Carolingian Empire,
Charlemagne (747/748–814) enacted the Capitulare de villis (ca 795) which
included also parsley. Up to the end of Middle Ages, parsley was mostly used as a
medical plant.
At present, it is grown in temperate and subtropical climate worldwide and pre-
dominantly used as aromatic plant for cooking and garnishing. In Germany parsley
13 Petroselinum crispum (Mill.) Nyman (Parsley)
438
is the most important spice plant, cultivated on more than 1800ha (Schmitz and
Pforte 2014). In the growing segment of potted spice plant production in Germany,
parsley ranked second (19%) behind basil (Ocimum basilicum L., 47%).
13.2.1 Plant Parts Used
All parts of the plant including leaves, stems and taproots are usable. Products from
harvested parsley leaves for seasoning of soups, sauces, dressings and meet dishes
and for garnishing are marketed dehydrated, frozen or as fresh green bunches. Since
some years pots with green parsley plants as pot herb are an increasing segment. Big
amount of parsley is used dehydrated for industrial processed foods. It is important
especially for soups and meat dishes.
Parsley roots are used fresh as vegetable and in soups. It is also often used for
avouring and to enhance the taste of meat in combination with celeriac, carrot and
leek. Dehydrated pieces of parsley root are used alone or as mix with other root
vegetables in a similar way as the fresh ones.
For the drying process horizontal belt dryers are used with temperatures starting
between 90°C and 120°C.With decreasing moisture of the processed leaves, the
drying temperature is reduced to 80°C to 60°C.The process nishes with 4–6%
residual moisture content (Hoppe etal. 2013). The relation of fresh weight to dry
weight for parsley leaves is about 7:1 and for parsley roots about 10:1. The drying
process for roots occurs with 40°C.The roots are sliced in small pieces before dry-
ing (Dachler and Pelzmann 1999).
The essential oil of parsley is applied in the food industry and as a fragrance in
perfumes and for cosmetics (Feldheim 1999).
Parsley has carminative, tonic and aperient effects, but it is mainly used for its
diuretic properties (Warncke 1994). It stimulates the appetite, has anti-inammatory
properties and has the ability to induce menstruation because it stimulates gentle
contractions of the uterus.
Traditionally, it has been used for atulent dyspepsia, colic, cystitis, dysuria,
bronchitis cough in the elderly, dysmenorrhoea, functional amenorrhoea, myalgia
and specically for atulent dyspepsia with intestinal colic.
The oral consumption of the essential oil (Petroselini aetheroleum) with high
content of apiole can lead to intoxications. Parsley is safe in normal food quantities
of green and dehydrated material (Hoppe etal. 2013).
The pharmaceutical use is not recommended by the European Medicines Agency
(EMA) because of the side effects especially in the use of essential seed oil. The
consumption of fresh or dehydrated leaves and root products is absolutely risk-free.
A urological phytopharmaceutical in Switzerland is Asparagus-P, lm-coated tab-
lets (active ingredients, Asparagi radicis pulvis, Petroselini herbae pulvis; company,
Phytaris Naturwissen GmbH). Parsley is used in homoeopathic pharmaceuticals
and in folk medicine.
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439
13.2.2 Cultivation
Parsley is a culture, which is grown on elds or greenhouses and plastic tunnels. It
has to be sown at beginning of the vegetation period. For the temperate climate of
Central Europe, this should be from beginning of March on. The plants form a
rosette of leaves and the rst harvest takes place 10–12weeks after start of cultiva-
tion. According to the weather conditions every 21–30days a harvest of the new
grown leaves can occur. For harvest the plants of at leaf type should reach a height
of 20–25cm. Varieties of the crispy leaf type are not so high. The cut should be
4–6cm above the ground (Hoppe etal. 2013).
For growing parsley, sandy and sandy clay soils are preferable with a relatively
high level of humus from 1 to 5% and a pH value from 6 to 7 (Dachler and Pelzmann
1999). Parsley is a cool season plant that grows optimally at temperatures between
7 and 16°C, while growth slows considerably below 7°C (Small 1997). The germi-
nation of parsley seeds is slow and can reach up to 26days under low temperature.
Germination is best at 12–15°C.
For parsley seeds, a pretreatment by soaking the seeds for 18hours in 20±2°C
warm water and subsequent re-drying at 30°C maximum temperature is useful.
Seed dormancy can be broken by stratifying in a refrigerator with around 5°C or
briey freezing.
Parsley leaves tolerate temperatures down to −6 to −8°C.Roots will survive the
winter with temperatures down to −10°C.However, alternating frosts can lead to
complete loss by rot. For seed production in the second year, it can be necessary to
uproot the root ends of growing season and to replant them in spring.
A crop rotation system that includes parsley should guarantee a growing inter-
mission of minimally 5years, better up to 7years between parsley but also to other
Apiaceae. Suitable preceding crops are species depressing weed well.
13.2.3 Valuable/Undesired Plant Secondary Compounds
Parsley is rich in antioxidants (especially in vitamin C, (CAS 50–81-7), in the avo-
noid luteolin (CAS 491–70-3), in vitamin A and in folic acid (vitamin B9, CAS
59–30-3). It contains essential oils in roots, leaves and fruits with clear differences
in amount and composition according to the used plant tissue (Hoppe etal. 2013).
The essential oil content of parsley roots of the taprooted form (P. crispum con-
var. radicosum) is with 0.1–0.3% lowest in relation to leaves (0.02–0.9%) and fruits
(1.0–6.0%). After steam distillation of taproots the main components of the essen-
tial oil are β-pinene (CAS 127-91-3, 21–40%), β-phellandrene (CAS 555-10-2,
7–14%), apiole (CAS 523-80-8, 15–42%) and myristicin (CAS 607-91-0, 5–15%).
Extracts of uneatable roots of P. crispum convar. crispum contain terpinolene (CAS
586-62-9, 7–43%), β-pinene (4–30%), apiole (20–57%), myristicin (4–21%), small
amounts of elemicin (CAS 487-11-6) and others (Franz and Glasl 1974; Gijbels
13 Petroselinum crispum (Mill.) Nyman (Parsley)
440
etal. 1985; Warncke 1994). Parsley roots contain avonoids (0.2–1.6%) with the
main component apiin (CAS 26544-34-3), furanocoumarins (to 0.1%) with oxypeu-
cedanin as main furanocoumarin (CAS 737-52-0), bergapten (CAS 484-20-8),
imperatorin (CAS 482-44-0), isopimpinellin (CAS 482-27-9) and methoxsalen
(CAS 298-81-7) (Baumann etal. 1988; Chaudhary etal. 1986), phthalides as ligust-
ilide (CAS 4431-01-0), senkyunolide (CAS 62006-39-7), butylphthalide (CAS
6066-49-5) (Gijbels etal. 1985), polyenes as falcarinol (CAS 21852-80-2) and fal-
carindiol (CAS 55297-87-5) (Bohlmann 1967; Nitz etal. 1990).
The essential oil content of leaves amounts 0.02–0.9%. A set of 219 accessions
which included the parsley set from Federal ex situ Gene Bank for Agricultural and
Horticultural Crop Species at the Leibniz Institute of Plant Genetics and Crop Plant
Research (IPK) at Gatersleben, Germany, was characterized for composition of
essential leaf oil (from the rst cut) by hydrodistillation and gas chromatography.
The main components were in 96 accessions 1,3,8-p-menthatriene (CAS
18368-95-1), apiole in 55 accessions, myristicin in 50 accessions, β-phellandrene in
12 accessions, α,p-dimethylstyrene (CAS 1195-32-0) in three accessions and myr-
cene (CAS 123-35-3) in another three accessions (Fig.13.2). Further components
are α-pinene (CAS 80-56-8) and β-pinene, α-phellandrene (CAS 99-83-2), p-cymene
(CAS 535-77-3), terpinolene, (E)-β-farnesene (CAS 18794-84-8), germacrene D
(CAS 37839-63-7), crispane (CAS 79803-28-4) and crispanone (CAS 11053-21-7)
(Spraul etal. 1992; Hoppe etal. 2013).
Minor components with small or very small levels are methyl 2-methylbutanoate
(CAS 868–57-5), 1-octen-3-one (CAS 4312-99-6), (Z)-1,5-octadien-3-one (CAS
65767-22-8), 2-(4-methylphenyl)propan-2-ol (CAS 1197-01-9), 2-isopropyl-3-
methoxypyrazine (CAS 25773-40-4), 2-sec-butyl-3-methoxypyrazine (CAS
24168-70-5), p-mentha-1,3,8-triene (CAS 18368-95-1), (Z)-6-decenal (CAS
105683-99-6), (E,E)-2,4-decadienal (CAS 25152-84-5), linalool (CAS 78-70-6),
citronellol (CAS 106-22-9) and β-ionone (CAS 14901-07-6) (PubChem 2019,
TGSC 2019, Bauermann etal. 1993, Franz and Glasl 1974, Lawrence 1981, 1990,
MacLeod etal. 1985, Pino etal. 1997, Porter 1989, Spraul etal. 1991, Warncke1994).
All these volatile components are involved in avour formation of parsley leaves.
Fig. 13.2 Absolute
numbers of main
component types in 219
accessions of the essential
leaf oil
F. M ar the
441
Parsley leaves contain avonoids (1.9–6.6%) with the main component apiin
(apigenin-7-O-(6-O-apiosyl)glucoside (CAS 86546–87-4)). Further components are
luteolin-7-apiosylglucoside, apigenin-7-glucoside, 6″-acetyl-apiin, isorhamnetin-
3,7-diglucoside and chrysoeriol-7-apiosylglucoside (CAS 33579–63-4) (Grisebach
and Bilhuber 1967; Nordstrom etal. 1953; Yoshikawa etal. 2000).
Furanocoumarins of parsley leaves reach up to 0.2%. The main components are
oxypeucedanin and bergapten (5-methoxypsoralen, CAS 484–20-8). Further fura-
nocoumarins in parsley are xanthotoxin (8-methoxypsoralen, CAS 298–81-7), pso-
ralen (CAS 66–97-7), imperatorin (CAS 482–44-0) and isopimpinellin (CAS
482–27-9) (Ashraf et al. 1980; Baumann et al. 1988; Chaudhary et al. 1986;
Teuscher and Lindequist 1994).
In fresh parsley leaves vitamin C (ascorbic acid, CAS 50–81-7) can reach
0.12–0.4% of fresh weight (Feldheim 1999; Manderfeld etal. 1997; Scheunert and
Theile 1952).
The essential oil content of fruits (Petroselini fructus) is 1.0–6.0% with the main
components apiole, myristicin and 1-allyl-2,3,4,5-tetramethoxybenzene (CAS
15361–99-6) (Hoppe etal. 2013). Because of the relatively high concentration of
essential oil, only fruits were used for essential oil production.
13.2.4 Main Production Areas
Parsley belongs to the most signicant medicinal and aromatic plants worldwide
regarding the area under cultivation and harvest size. For international export pars-
ley production of Egypt is the most important for dehydrated parsley as well as for
green bunches and frozen spice. Uzbekistan is important for production and export
to other Asian countries. However, parsley growing is also important in India,
Malaysia, Indonesia, China and the Philippines. For America Canada is an impor-
tant producer of dehydrated parsley akes. The United States with production con-
centration in California, New Jersey, Florida and Texas and smaller volumes from
Ohio, Massachusetts and other states is a signicant producer and a signicant mar-
ket for fresh and dehydrated parsley (Small 1997).
For the member states of European Union, around 5.000ha are estimated.
Production of dehydrated parsley is important in Germany, France, the
Netherlands and Poland. Production of green bunches and frozen spice food is
important in France, Belgium, the Netherlands, Great Britain, Germany and
Poland. Production of potted plants comes from the Netherlands, Germany,
Belgium, France, Great Britain and Poland (Junghanns 2018). For turnip-rooted
parsley the United Kingdom is an important production area. Countries of Central
and South Eastern Europe Slovakia, Romania and Bulgaria have tradition in
growing and using turnip-rooted parsley. Many accessions in gene bank come
from this region.
13 Petroselinum crispum (Mill.) Nyman (Parsley)
442
13.2.5 Economical Valuation/Parameters
Under the conditions of Germany the yield of dry mass of rst cut ranges from 800
to 900kg/ha. For the second cut 1.0–1.1t/ha are possible. All in all up to 5 cuts per
year and up to 5.3t/ha dry mass per year are possible. The importance of early sow-
ing was demonstrated by Kienast (Hoppe etal. 2013). They reached 2012 6.1t/ha
and year dry mass yield by sowing in February, 12 kg/ha seed rate and distance
between rows of 12–13cm.
Essential oil produced from parsley is of lower value than fresh and dried prod-
ucts. Verlet (1993) estimated the annual world value of parsley essential oil from
seed to be $600,000.00 (US), while that of “parsley herb” essential oil amounted to
an additional $360,000.00 annually.
13.2.6 Pests andDiseases
The cultivation of parsley on elds or in greenhouses requires a monitoring of phy-
topathological situation of this crop.
For Central Europe behind pests especially fungi, viruses and bacteria are of
signicance. The requirement of products without phytopathological damages
stands in conict with the interest of consumers in goods produced nearly without
or without chemical pesticides and the decreasing number of approved active sub-
stances. The best way to produce parsley without residuals is to use resistances to
the pathogens. For breeding projects, information about the biology of the patho-
gens is necessary. Also evaluation data for sources of resistance are essential.
Economically important diseases are Septoria blight caused by Septoria
petroselini (Lib.) Desm., downy mildew (Plasmopara petroselini Săvul. &
O.Săvul.), powdery mildew (Erysiphe heraclei DC. ex Saint-Aman) and Alternaria
leaf blight caused by Alternaria radicina Meier, Drechsler & Eddy.
Septoria blight (S. petroselini) is worldwide one of the most important pathogens for
parsley. It is seed-borne (Tahvonen 1978). Pycnidia are xed permanently within the
pericarp of the schizocarpic fruit, containing pycniospores for mass infection (Ferri 1969).
Lesions of Septoria blight are round or angular and greyish brown, clearly bor-
dered by a darker brown margin to the surrounding tissue, and have small or no
chlorotic zones (Fig.13.3). Spores containing black pycnidia appear within lesions
(Meyer etal. 2010; Marthe and Scholze 1996; Cerkauskas and Uyenaka 1990).
Plasmopara petroselini was separated from Plasmopara nivea by Săvulescu and
Săvulescu (1951).
Downy mildew on parsley was found several times in Germany at the end of the
nineteenth and the rst half of the twentieth century (Brandenburger and Hagedorn
2006). Together with Septoria blight, they are currently the most relevant pathogens.
The downy mildew causing fungus P. petroselini induces faint chlorotic spots on
the upper surfaces of the leaves as rst symptoms. On the corresponding lower sur-
faces, sporangiophores and white-to-greyish white mycelium develop (Fig.13.4).
Lesions grow fast and parts of the leaf or whole leaves rot (Amein etal. 2006;
F. M ar the
443
Crepel and Inghelbrecht 2003). Spores are small, oval and hyaline. Downy mildew
is an important pathogen in the production of potted parsley. Infection spreads very
rapidly in greenhouse or plastic tunnel especially in the period of shorter days. The
infected crop develops a typical smell.
Powdery mildew (E. heraclei) on parsley is a worldwide signicant pathogen
(Marthe etal. 2003; Scholze etal. 1996). The economic importance of E. heraclei
Fig. 13.3 Septoria blight
on parsley; (a) lesions; (b)
lesions with black pycnidia
Fig. 13.4 Downy mildew on curled leaf parsley; (a) plants with necrotic leaves; (b) lower leaf
surface with necrosis and sporangiophores
13 Petroselinum crispum (Mill.) Nyman (Parsley)
444
for production under eld conditions in Germany is lower than S. petroselini and
P. petroselini. The rapidly increasing production of parsley pots around the year is
much more affected by E. heraclei. Powdery mildew attacks also other members of
Apiaceae family. This pathogen may occur early in the season and cause losses in
quality of the market value of the product. E. heraclei on parsley has an increasing
range worldwide. The fungus was rst described in Brazil: Sao Paulo, E. heraclei,
introduced through contaminated seeds (Rosa etal. 2008).
In the USA, the disease was several times observed and described (Raid etal.
2007; Glawe etal. 2005; Koike and Saenz 1994). In Europe, powdery mildew has
been an important pathogen for a long time.
The identication is easy because of the mycelia growing on the leaf surface
(Fig.13.5). Haustoria of oval shape establish in epidermis cells for absorbing nutri-
ents (Fig.13.6). Infected leaves show chlorosis and crinkling (Marthe etal. 2003;
Koike and Saenz 1994).
Alternaria radicina Meier, Drechsler & Eddy (syn. Stemphylium radicinum
[Meier, Drechler & Eddy] Neerg.) causes Alternaria leaf blight and damping off of
seedlings (Gärber etal. 2007; Marthe and Scholze 2006; Nawrocki 2005; Marthe
etal. 2003; Nowicki 1997; Gärber and Ulbrich 1996).
Infestation by A. radicina causes important damages by leaf spots (Meyer etal.
2010) and loss of quality depending on annual weather conditions. Alternaria spe-
cies are seed born on parsley. The conidia adhere together with mycelium external
on the seeds and so they are good diagnosable.
Infestation starts from older leaves with brown-coloured small round lesions. At
this stage the differentiation of lesions to early infestation by S. petroselini is nearly
impossible. But with progression of the disease, characteristic lesions will develop.
The Alternaria lesions expend rapidly usually from margin of leaf and are shaped
irregularly. Inside the lesions, concentric zones can be visible with moderately
graded colours. The transitions to non-necrotic area of leaf are uid conditioned by
a chlorotic zone (Fig.13.7). Inside the lesions and successional on the whole dieback
Fig. 13.5 Powdery
mildew on parsley; (a)
plants with ecto-mycelium;
(b) leaf with mycelium and
mass production of asexual
hyaline spores
F. M ar the
445
leaves grey to black conidia are built terminally at the end of branched mycelia. This
is the reason for the characteristic black colour of infested parts of the plants.
In nearly in all cases associated with specialized A. radicina the species Alternaria
alternata [Fr.] Keissler (synonym Alternaria tenuis C.G.Nees.) followed as sapro-
phyte. This species is not specialized to species or plant families and widespread
through climatic zones. The conidiospores evolve as chains by constriction from
mycelia. A. alternata is also able to set very small lesions on parsley leaves. For this
a longer period with high air moisture and around 20°C are necessary. However, the
damages are not of economic importance.
Fig. 13.6 Powdery
mildew on parsley,
haustoria formed in
epidermis cells; coloured
with Cotton Blue
Fig. 13.7 Alternaria leaf
blight on parsley; (a)
plants with necrotic leaves,
(b) leaf with necrosis and
chlorotic zones
13 Petroselinum crispum (Mill.) Nyman (Parsley)
446
The generalist Sclerotinia sclerotiorum (Lib.) de Bary causing Sclerotinia rot
(Nowicki 1997; Scholze etal. 1996) are also relevant phytopathogen for parsley. It
is easy to identify by the more or less big black sclerotia embedded in cotton-like
mycelia. Attacked plants are destroyed by macerating of the stalk basis.
There are more pathogenic fungi damaging parsley as Fusarium wilt (Fusarium
oxysporum Schltdl.) (Nawrocki 2005; Scholze et al. 1996), Pythium root rot
(Pythium spp.) (Tsuchida etal. 2018; Petkowski etal. 2013; Gärber and Borchers
2000; McCracken 1984) and Rhizoctonia root rot (Rhizoctonia solani J.G.Kühn)
(Nawrocki 2005; Monnet and Thibault 2002; Uematsu etal. 1993).
Viruses of economic impact on parsley are the widespread Celery mosaic virus
(CeMV), the Apium virus Y (ApVY) and a virus complex probably from the Carrot
red leaf virus, (CtRLV), the parsley-specic Carrot mottle virus (CMoV) and Carrot
mottle mimic virus (CMoMV). In addition, the Apium virus Y and the Carrot yellow
leaf virus (CYLV) are possibly involved in this virus complex (ICTV 2019; Hoppe
etal. 2013; Meyer etal. 2010).
Bacteria can cause leaf spots and macerations, especially by Pseudomonas spp.
(Reintke etal. 2016; Bull etal. 2011).
13.3 Breeding
Breeding Objectives
For optimization of parsley lines by using methods of plant breeding many require-
ments exist. For breeding success as well as for economic result the choice of breed-
ing aims is of particular importance.
For many crops, the group of yield parameters is of big or biggest interest. That
is also true for parsley. Breeding objectives of this group for leaf production are size
of yield, amount of marketable yield, essential oil content, fast regrowth after cut-
ting, colour of leaves, crimping of the leaf blade tissue between the secondary veins,
relation of leaves to stalk, height and density of plants.
Breeding objectives for turnip-rooted parsley are yield of taproots, length and
diameter of the taproot and a smooth taproot.
For many of these characters the International Union for the Protection of New
Varieties of Plants (UPOV) has exact descriptors under UPOV code: PETRO_CRI
(UPOV 2005, Table13.1, Figs.13.8 and 13.9).
A considerable number of breeding aims exist for resistances to pathogens and for
stress tolerance. Usable resistances can be deciding for the qualication to be culti-
vated. This is evident for ecological cultivation but is also true for controlled cultiva-
tion. The availability of active components for plant protection is restricted. In the near
future the number of such substances for pest management will decrease drastically.
In extensive tests for resistance to Septoria blight caused by S. petroselini no
resistant variety could be found. For these tests it was necessary to develop a method
for cultivating the fungus on articial medium and a method for testing the resis-
tance in climate chamber (Marthe and Scholze 1996). The pycniospores necessary
for articial inoculation were produced on articial vegetable juice medium in petri
F. M ar the
447
dishes under sterile conditions. Concentration of spores in a water suspension was
approximately 106/ml. Young plants were sprayed with 3ml of the suspension per
plant. After inoculation plants were incubated under controlled conditions at tem-
peratures of 18–20°C and a day/night light regime of 16/8h. For good infection
results it is necessary to provide more than 95% humidity for at least 4 days after
inoculation. After an incubation time of 21days, when susceptible leaves are show-
ing lesions, disease assessment was carried out. No variety or gene bank accession
was found free of symptoms for all single plants. However, clear differences in
Table 13.1 Characteristics and examples from varieties of parsley, including also turnip-rooted
parsley
Number an footnotes Characteristic
1. (∗) Plant: height
2. Plant: width
3. (∗) Plant: density of foliage
4. Plant: number of leaves
5. Leaf: attitude
6. (∗) Leaf blade: curling
7. (∗), (+) Leaf blade: intensity of curling
8. Only varieties with leaf blade curling
Plant: appearance of surface of canopy
9. Only varieties with leaf blade curling
Leaf blade: upward reexing of lobes
10. (∗) Leaf blade: length
11. (∗), (+) Leaf blade: width
12. Leaf blade: ratio length/width
13. (∗) Leaf blade: intensity of green colour
14. (+) Leaet: shape
15. (+) Leaf blade: distance between rst and second pair of leaets
16. Leaet: undulation of margin
17. (+) Petiole: length
18. (+) Petiole: thickness
19. (+) Petiole: anthocyanin coloration
20. (∗) Root: thickening of main root
21. (∗) Only root parsley varieties
Root: length
22. (∗) Only root parsley varieties
Root: width
23. (∗)Only root parsley varieties
Root: ratio length/width
24. Only root parsley varieties
Root: branching
From UPOV (2005), simplied
(∗): Characteristics included in the Test Guidelines which are important for the international har-
monization of variety descriptions and should always be examined for DUS and included in the
variety description by all members of the Union, except when the state of expression of a preceding
characteristic or regional environmental conditions render this inappropriate
(+): See Fig.13.9
13 Petroselinum crispum (Mill.) Nyman (Parsley)
448
Fig. 13.8 Classication of the curling intensity of the leaf blade: (a) weak, 3; (b) medium, (c) 5;
strong, 7; (d) very strong, 9. (From UPOV 2005)
Fig. 13.9 Instruction for
measuring of characters
listed in Table13.1: (10)
leaf blade, length; (11) leaf
blade, width; (15) leaf
blade, distance between
rst and second pair of
leaets; (17) petiole,
length; (18) petiole,
thickness. (From UPOV
2005)
F. M ar the
449
reaction to the pathogen have been found between the accessions of germplasm
material tested in climate chamber. This type of resistance is characterized by popu-
lations including plants with low level of symptoms. In resistant populations, rst
symptoms appear later and infestation is lower in comparison to highly susceptible
varieties and accessions. From resistant populations, plants free or nearly free of
symptoms were selected and self-pollinated to increase homozygosity.
In a resistance test under natural infection pressure 220 accessions were scored.
The rating scale consists of level 0 (without any lesion) to 9 (plants die by action of
pathogen). Level 9 has not be observed in our sample set. Scoring rates 0 and 1 are
classied as ‘free or nearly free of symptoms’, 3–5 are ‘moderately susceptible’ and
7 and 8 are ‘highly susceptible’. Only one accession was free of symptoms. The
results generate a distribution similar to a Gaussian curve (Fig.13.10). This charac-
terizes the quantitative action of the resistance (Marthe etal. 2013). Crossing exper-
iments for characterization of resistance inheritance indicate an oligogenic situation
(Bruchmüller 2013).
Since the beginning of the twenty rst century downy mildew caused by
P. petroselini is as an economically important pathogen. Therefore, the availability
of resistances is of high interest. Leinhos etal. (2012) developed a method for test-
ing downy mildew resistance. For inoculation a spore suspension of 1 x 105 sporan-
gia/ml in water was used. The temperature for infection is from 4°C to 15°C with
an optimum of around 10°C.The temperature is correlated with duration of leaf
wetness. For a high infection rate 24hours (4°C) to 4hours (15°C) are necessary.
The latency period was calculated with 8days (14–24 °Cday temperature, 3°C
night temperature and 77% relative air moisture). Twelve days after infection, the
area of infested leaves and sporulation reached its maximum.
First results from tests with inoculation indicate that variety ‘Felicia’ was free of
symptoms out of nine varieties tested (Krauthausen and Leinhos 2007).
Fig. 13.10 Number of parsley accessions scored for lesions by Septoria petroselini under natural
infection. Scoring rates 0, 1: free or nearly free of symptoms; 3, 5: moderately susceptible; 7, 8:
highly susceptible. (From Marthe etal. 2013)
13 Petroselinum crispum (Mill.) Nyman (Parsley)
450
In a test under natural infection Marthe etal. (2013) found 73 accessions free or
nearly free of symptoms under 219 accessions and varieties. The results generate a
bimodal distribution of accessions free of symptoms and susceptible accessions
(Fig.13.11). This is an indication for existence of qualitative resistance. The acces-
sions free of symptoms of P. petroselini together with variety ‘Felicia’ tested free of
symptoms are rst candidates for intraspecic resistance in parsley. These candidates
should be tested again for level of resistance. The accessions free of symptoms also
open up the possibility to look for candidates for tests of race differences in
P. petroselini. From many other downy mildew varieties a high number of different
races are known (Marthe etal. 2013).
Powdery mildew caused by E. heraclei is also signicant for parsley production,
especially for production of potted plants and fresh green leaves. But also eld cul-
tures can be attacked by powdery mildew depending on weather conditions.
Powdery mildew is an obligate biotrophic pathogen. The cultivation on articial
medium is an unsolved problem. The pathogen has to be cultivated on living plants
for mass production of spores and storage of isolates. For resistance test, Marx and
Gärber (2014) developed a method for inoculation. The germination of the conidia
correlates with temperature. At all tested temperatures from 6°C to 30°C conidia
are germinable. The highest level of germination with 80% was from 15°C to
20°C.Lower and higher temperature reduced the level of conidia germination. The
temperature inuences also the length of incubation period. The shortest period was
7days at the temperature of 25°C.Lower temperatures (10°C, 18days) and higher
temperatures (30°C, 12days) prolonged the incubation period. However, the level
of air moisture (30, 50, 70% at 25°C) has no inuence on the length of incuba-
tion period.
70
51
22
65
Plasmopara petroselini
20
57
4
60
50
40
30
20
10
0
013
Scoring rate
Number of accessions
578
Fig. 13.11 Number of parsley accessions with scoring rates 0, 1: free or nearly free of symptoms;
3, 5: moderately susceptible; 7, 8: highly susceptible to natural infection by Plasmopara petroselini.
(From Marthe etal. 2013)
F. M ar the
451
For sporulation the level of air moisture has inuence. The number of produced
conidia increases with higher air moisture. The maximum sporulation was reached
at 70% air moisture. At temperatures from 20°C to 25°C level of sporulation was
higher than on 15°C (Marx and Gärber 2012).
Although E. heraclei has been a signicant pathogen in Europe for a long period,
no results are available of extensive evaluation for resistant and susceptible acces-
sions. Two evaluations of gene bank accessions were made under natural infection
(Marthe etal. 2013; Marthe etal. 2003). From this, it was concluded that resistances
exist in the tested set. Before nal judgement of resistance, the candidates for resis-
tance have to be tested under controlled conditions with inoculation.
Marx and Gärber (2012) tested 11 isolates of powdery mildew from parsley (10)
and dill (Anethum graveolens L., 1) on two varieties of parsley. The scoring 15days
after inoculation generated nearly 100% of infected leaf surface on half of the iso-
lates. The other half infected only 25% of leaf surface.
These results demonstrate the necessity for testing race specicity of different
isolates of E. heraclei because for many other varieties of species of powdery mil-
dew high numbers of different races are known. Variability in response to powdery
mildew was evident offering possibilities to develop resistant cultivars.
Alternaria leaf blight causes leaf necrosis, which results immediately in a loss of
quality. Under Central European conditions it can be found in nearly each year, but
major damages depend on weather conditions and are not likely to occur frequently
or regularly. For evaluation of collections including gene bank accessions for
sources of resistance a technique was adapted for tests under controlled conditions
in a climate chamber. A. radicina and A. alternata can be cultivated on articial
medium. On malt extract agar (25ml/l, 23g/l agar-agar, 0.25mg/l chlorampheni-
col) monosporic isolates develop under sterile conditions at room temperature large
amount of spores. Suspension of conidiospores in water (1×106 conidia/ml) were
sprayed on six-week-old plants (3–4ml/plant). Inoculated plants were cultivated in
a climate chamber with saturated air moister, temperature of 19°C and light of
16hours. Disease assessment was carried out 7days after inoculation and a second
time 14days after inoculation by estimating the lesions in relation to leaf surface
(Marthe and Scholze 2006).
The results of different tests with a set of varieties and gene bank accessions
indicate clearly resistances in the tested set. It was possible to differentiate two
material groups that differ in the level of adaptation to temperate European growing
conditions. The number of resistant varieties from the European temperate zone was
higher than accessions from other zones. There are resistances to A. radicina inside
the European breeding material. However, the control of resistance in the breeding
process is recommended (Marthe etal. 2003).
From the use as spice plant, signicant quality parameters are of importance.
The most important is the shelf life. Because of relatively fast green colour loss
of parsley bunches, the production of potted parsley plants gained in importance.
In the market consumers prefer minimally processed vegetables and a broad
spectrum of fresh aromatic plants. For this the shelf life characteristics are of impor-
tance. Parsley has antibiotic acting secondary metabolites. Nevertheless, different
13 Petroselinum crispum (Mill.) Nyman (Parsley)
452
microbes occur on and inside the leaves. Unhygienic contaminations in the produc-
tion process by bacteria harmful to consumers can only be solved by hygienic stan-
dards (Santos etal. 2014). For this the post-harvest processing as fast drying starting
with temperature of 90°C and higher and also the pressurized CO2 disinfestation
process has proven its reliability.
The complexes of smell and taste are also important. The optimization of this
breeding aims should be handled with respect to resistance or repellent action of
some sensory effective substances, which are also included in resistance responses.
Parsley is known for both an outstanding and unique avour and bioactive sec-
ondary metabolites (Cazzola etal. 2011). The sensory quality as well as the compo-
sition of volatile compounds of freshly harvested parsley, dried parsley and essential
oil has been studied in the past (Masanetz and Grosch 1998a, 1998b; Jung etal.
1992; Whiteld and Last 1991; MacLeod etal. 1985). A total of around 80 volatiles
have been identied, of which a smaller number of 17 odorants show a relatively
high aroma impact. The avour of freshly harvested and cut parsley leaves is caused
mainly by p-mentha-1,3,8-triene, myrcene, 2-sec-butyl-3-methoxypyrazine, myris-
ticin, linalool, (Z)-6-decenal and (Z)-3-hexenal (Ulrich etal. 2011).
Hoberg etal. (2007) developed a method for the evaluation of sensory features
and their variability by a panel of testers. Every test person tastes the standard vari-
ety ‘Grüne Perle’ and collects descriptive, objective attributes for the categories.
The resulting method includes 29 suitable sensory attributes for taste, odour, retro-
nasal odour and mouth feeling. In addition, the estimated hedonic acceptability was
noticed by each tester. The sensory method was adapted to breeding and does not
focus on characterization of substances and their impact on sensorially overall
impression. It enables evaluation of varieties and gene bank accessions as well as
the monitoring over the breeding process. The sensorial characteristics and the con-
tent and composition of metabolites of most accessions conserved in gene banks are
not characterized. In the set tested by Hoberg etal. (2007) it was found that the year
of harvesting, the resistance against Septoria petroselini (Lib.) Desm. and above all
the habitus have a signicant effect on the majority of the sensorial parameters.
However, so far the interrelations between aroma compounds and their function-
ality in resistance mechanisms are widely unknown, not only in parsley. The results
show that no simple and straight correlation exists between sensorial quality and
resistance properties. The recognized variability of the sensory features and the
described method for their objective evaluation offer good chances to breed well
tasting parsley with resistances to pathogens in the future.
The group of breeding aims resulting from requirements of pharmacopoeia is for
the factual scope of European Pharmacopoeia application not of special interest. A
monography for parsley does not exist in Ph.Eur. 9 (2017) besides the general
requirements of the monograph herbal drugs. The German Homoeopathic
Pharmacopoeia (Homöopathische Arzneibuch HAB 2018) included Petroselinum
crispum convar. crispum for use as fresh whole plant at stage of owering begin
with no requirements for metabolites.
F. M ar the
453
13.3.1 Flower andPollination Biology
Flowers and Flowering: Pollination
After the rst winter, the stalk of biennial parsley will shoot immediately. The leaves
of the ower stalk are smaller and pinnation is simpler. Flowering plants reach a
height of 60–120cm. At the end of the stalk umbels of different ranks are formed.
Inside the umbels, a time gradient exists with the oldest partial inorescences at the
external circle and the younger unripen owers in the internal circles. Such a gradi-
ent exists also inside the partial inorescences, which group owers arising from the
same point. Each ower has ve petals. The colour of petals varies between acces-
sions from green to white (Fig.13.12).
Protandry is common in the family Apiaceae and occurs in parsley as well. The
ve anthers start anthesis around 5 days before the two stigmata are receptive to
pollen (Fig.13.13). At the time of receptiveness for pollination, the receptacle nec-
taries produce a sugar-rich liquid for rewarding pollinating insects. At this time, the
ower gets shiny and the stigmatic papillae reach the length of ower radius. In a
ower two carpels are fused to a bicarpellate pistil from which the schizocarpic
fruits develop (Fig.13.14). The mature fruits can break into two mericarps of one
single fruit (commonly called ‘seed’) each.
Male Sterility
Male sterility is not seldom in family Apiaceae. The model plant for this is carrot
(Daucus carota L.) where cytoplasmic male sterility (cms) is used in practical
breeding. Two types of male sterility exist in carrot: the brown anther type and the
petaloid type. The molecular mechanisms for cms in carrot are under investigation
(Tan et al. 2018; Szklarczyk et al. 2014; Robison and Wolyn 2006). For celery
(Apium graveolens L.) male sterility was found and characterized (Quiros etal.
1986). For fennel (Foeniculum vulgare MILL.) the existence and use of male steril-
ity is documented (Palumbo etal. 2018; Pank etal. 2007). For parsley no source for
male sterility or cytoplasmic male sterility was found. In own investigations on
about 200 accessions and varieties no male sterile plants were detected.
Fig. 13.12 Flowers of parsley; left, green petals; right, white petals
13 Petroselinum crispum (Mill.) Nyman (Parsley)
454
Hybridizations
Crossing technique is a prerequisite to obtain genetic recombination at the begin-
ning of the breeding process or genetic analyses. For hybridization, emasculation of
an umbel has to be performed. The anthers of the earliest third of the owers should
be fully developed. At this time, the ower is not ready for pollen reception. All
older owers with receptive papillae have to be removed manually by a tweezer. For
this a head loupe generates a stereoscopic image. Furthermore, the younger owers
and buds have to be removed by a tweezer because their anthers would develop pol-
len. The remaining owers with unripe papillae have to be ushed out by a strong
water jet to remove all pollen. At the end, the emasculated umbel has to be con-
Fig. 13.13 Inorescence of parsley; left, (double) umbel; right, ower with the two immature and
non-receptive stigmata
Fig. 13.14 Inorescence of parsley with almost ripe schizocarpic fruits
F. M ar the
455
trolled for remaining anthers and then isolated by a bag. The material of the bag can
be a translucent plastic lm with micropores. Three to ve days later the pollination
has to be performed by touching the papillae with owering umbels from the male
crossing partner. After pollination the emasculated umbel has to be isolated again by
a plastic bag. About 1 week later the fruit growth should start and the bag can be
removed.
13.3.2 Propagation Strategies
Generative Propagation
After winter the roots of the vegetative growing period are used for seed production.
The harvest takes place when the rst seeds are ripe. Selective harvest of oldest
umbels decreases the loss of fully ripened seeds but is labour intensive.
Vegetative Propagation
There are no strategies for vegetative propagation in parsley.
13.3.3 Breeding Methods Applied
Conventional Breeding
The breeding of parsley started with selections from land races. An example is the
curled parsley ‘Mooskrause’ from the nineteenth century, which was divided into
‘Mooskrause 1’, ‘Mooskrause 2’ and ‘Mooskrause 3’.
Different breeders started programs for maintenance of the variety. This resulted
in differences between the selections ‘Mooskrause 2’ (Germany), ‘Moskrul 2’ (the
Netherlands), ‘Moss Curled 2’ (Great Britain), ‘Nain Frisé mousse’ (France) or
‘Nano ricciuto 2’ (Italy). From this so-called umbrella variety many new varieties
were developed by selection.
Other examples are the at-leaf parsley ‘Amsderdamse Snij’, ‘Einfache Schnitt
3 Typ Hamburger Schnitt’ and the turnip-rooted parsley ‘Halblange- Berliner’.
Interspecic crosses between parsley and celery (Apium graveolens L.) has been
successfully implemented since the beginning of the twentieth century (Becker-
Dillingen 1926, Becker 1943/44, Skiebe cited from Becker 1962). These sexual
crosses were successful only with celery as female. Later on, also reciprocal crosses
were done (Madjarova etal. 1973; Madjarova and Bubarova 1978; Madjarova 1978;
Honma and Lacy 1980; Lacy and Honma 1981). The aim of these crosses was to get
higher yielding bastards but primarily to transfer resistance against Septoria blight
into the bastard. Celery is resistant to Septoria petroselini and parsley to Septoria
apiicola Speg. Some of the produced bastards were more tolerant to the S. apiicola
(Madjarova etal. 1973; Honma and Lacy 1980) but they report the resistance as
13 Petroselinum crispum (Mill.) Nyman (Parsley)
456
polygenetic and recessive. The resistances were instable and had no practical impact
on breeding resistant varieties (Ochoa and Quiros 1989). The variety ‘Festival 68’
from Bulgaria was developed from an interspecic bastard of parsley and celery
(Madjarova and Bubarova 1978) but is susceptible to S. petroselini.
Hybrid Breeding
In the species P. crispum a tremendous variability is present. Also groups of differ-
ent origin and consequently high phylogenetically distance exist. This is an impor-
tant prerequisite for hybrid variety breeding. However, the lack of male sterility
elements and no information on the intensity of hybrid effect prevented hybrid
breeding.
Molecular Tools
The small budgets for molecular analyses in parsley as part of the medicinal and
aromatic plants causes the lack of using molecular techniques in the past. With the
development of new methods especially in next-generation sequencing and a price
drop, it is possible to use this approach also in minor crops. From genome-based
sequencing (GBS) AP2/ERF transcription factors from the transcriptome sequence
under different abiotic stresses were detected (Li etal. 2014). Such data open the
chance to use molecular methods for intensive parsley research.
Genome-wide association study (GWAS) resulted in not veried candidates for
marker associations with curling of leaves, formation of turnip roots and lignica-
tion of the root (Bruchmüller 2013). The evaluation of different genotypes included
also the analysis of sensorial characteristics in relation to the nontargeted volatile
patterns (headspace-SPME-GC) and resistance to Septoria petroselini. The more
resistant genotypes are characterized by several negative sensory characteristics. In
contrast, the contents of some volatile compounds correlate highly and signicantly
either with resistance (e.g. hexanal and alpha-copaene) or with susceptibility (e.g.
p-menthenol). Some of these compounds with very strong correlation to resistance
are still unidentied and are presumed to act as resistance markers (Ulrich etal.
2011; Hoberg etal. 2007).
Phylogenetic distances have been calculated by using of RAPD and ISSR mark-
ers (Domblides etal. 2010). The analysis of 32 populations, mostly varieties, but
also seven accessions of Russian gene bank VIR, St. Petersburg, describe consider-
able distances between accessions.
13.3.4 Breeding Results Achieved/Economical Transfer
(Registered Cultivars/Patents, Trial Results)
There exist a high number of varieties. But the high number of local races or land
races is of special interest. Table 13.2 contains the actual registered varieties for
European Union.
F. M ar the
Variety name Country maintainers Synonym
Used plant
part (1, 2)
36004 RZ a NL 108 (Rijk Zwaan Zaadteelt en
Zaadhandel BV)
(1) Leaf2
36504 RZ a NL 108 (Rijk Zwaan Zaadteelt en
Zaadhandel BV)
(1) Leaf2
A grosse racine
gros hâtif
b FR x Root1
Afrodite b DK 57 (Hild Samen GmbH) Leaf1
Alba a CZ 1495 (Moravoseed CZ a.s.), a
HU 102445 (Moravoseed s.r.o.)
Root1
Alto a FR 8067 (Vilmorin) Leaf1
Amsterdamse Snij b NL x Fijne Snij NL Leaf1
Andrei a RO 1031 (Staţiunea de Cercetare-
Dezvoltare pentru Legumicultură
Bacău)
Root2
Arat b NL 8 (Bejo Zaden BV) Root1
Arctica a NL 8 (Bejo Zaden BV) H Root1
Argon a NL 26 (Enza Zaden Seed
Operations BV)
Leaf1
Aroma a NL 8 (Bejo Zaden BV) Root1
Arsem a RO 1072 (S.C.Unisem
S.A.Bucureşti)
Root2
Astra a CZ 1495 (Moravoseed CZ a.s.) Leaf1
Atika a CZ 239 (SEMO a.s.) Root2
Berliner = Halblange Root1
Berlinova a DE 9989 (Satimex Quedlinburg
GmbH)
Root1
Bravour b NL x, b UK 176 (J.E.Ohlsens
Enke A/S)
Leaf1
Catalogno = Gigante di Napoli Leaf1
Champion b UK 189 (A.L.Tozer Ltd.) Leaf1
Commun 2
Comun 2
Comune 2
Plain Leaved
(Sheeps) 2
Comum 2
b FR x
b ES x
b IT 125 (SAIS Società agricola
italiana sementi)
b UK 38 (E.W.King & Co. Ltd.)
De hoja lisa 2 ES
Plain or single 2 IT
Simple 2 FR
Toscano 2 IT
Leaf1
Comum 2 = Commun 2 Leaf1
Comun 2 = Commun 2 Leaf1
Comun 3
Gewone Snij 3
b ES 3051 (Royal Sluis)
b NL 78 (SVS Holland BV)
De hoja lisa 3 ES
Einfache Schnitt 3 NL
Leaf1
Comune 2 = Commun 2 Leaf1
Cukrowa a PL 67 (Krakowska Hodowla i
Nasiennictwo Ogrodnicze POLAN
sp. z o.o.)
Root1
Curlina Leaf1
(continued)
Table 13.2 Plant variety catalogue for parsley (Petroselinum crispum, H-25) of plant variety
database– European Commission (2019)
458
Table 13.2 (continued)
Variety name Country maintainers Synonym
Used plant
part (1, 2)
Danubiu a RO 1072 (S.C.Unisem
S.A.Bucureşti)
Leaf2
Darki b DK 57 (Hild Samen GmbH), b
UK 176 (J.E.Ohlsens Enke A/S)
Leaf1
Darklett a DE 2549 (Hild Samen GmbH) Leaf1
De hoja lisa 2 = Commun 2 Leaf1
De hoja lisa 3 = Comun 3 Leaf1
Doble rizado = Frisé vert foncé Leaf1
Dobra a CZ 256 (Seva-Flora s.r.o.) Root2
Domaći lišćar b HR 177 (Podravka d.d.) Leaf2
Efez a CZ 239 (SEMO a.s.) Root2
Einfache Schnitt 3 = Comun 3 Leaf1
Extra triple curled
2
= Mooskrause 2 Leaf1
Favorit b NL 8 (Bejo Zaden BV) Leaf1
Félhosszú = Halblange Root1
Fest a CZ 1495 (Moravoseed CZ a.s.) Leaf1
Festival 68 a CZ 1495 (Moravoseed CZ a.s.), a
PL x
Leaf1
Fidelio a NL 26 (Enza Zaden Seed
Operations BV)
Leaf1
Fijne Snij = Amsterdamse Snij Leaf1
Francesa Frisada = Frisé vert foncé Leaf1
Frisé vert foncé
Gekrulde
Donkergroene
Doble rizado
Francesa
Frisada
b FR x, b NL 65 (Pieterpikzonen
BV)
b ES x
Rizado verde oscuro ES Leaf1
Gala a PL 356 (Hortag Seed Co.) Leaf2
Gazela a PL 854 (Vera-Agra sp. z.o.o.) Root2
Gekrulde
Donkergroene
= Frisé vert foncé Leaf1
Gekrulde b NL x Leaf1
Gewone Snij 3 = Comun 3 Leaf1
Gigante catalogno = Gigante di Napoli Leaf1
Gigante d’Italia = Gigante di Napoli Leaf1
Gigante d’Italia = Gigante di Napoli Leaf1
Gigante di
Chioggia
= Gigante di Napoli Leaf1
Gigante di Napoli
Gigante d’Italia
b IT x
a DE 2549 (Hild Samen GmbH)
Catalogno IT
Gigante catalogno IT
Gigante d’Italia IT
Gigante di Chioggia IT
Verde scuro d’Italia IT
Leaf1
(continued)
F. M ar the
459
Variety name Country maintainers Synonym
Used plant
part (1, 2)
Grüne Perle a DE 86 (Karl und Walter Hild) Leaf1
Halblange
Halvlang
Berliner
Halange
Félhosszú
a CZ x, a DE x, a PL x, a SK x
b NL x
a HU 151508 (ZKI
Zöldségtermesztési Kutató Intézet
Zrt.)
Root1
Halange = Halblange Root1
Halvlang = Halblange Root1
Hamburska a PL 356 (Hortag Seed Co.) Root2
Hanácká a CZ x, a SK 250 (Zelseed spol. s
r.o.)
Root1
Ines a DE 7092 (GHG Saaten GmbH) Leaf1
Jadran a CZ 239 (SEMO a.s.) Root1
Jagienka a PL 92 (PlantiCo Hodowla i
Nasiennictwo Ogrodnicze Zielonki
sp. z o.o.)
Root2
Junák a SK 432 (P.K.SEM spol. s.r.o.) Leaf2
Kadeřavá a CZ x Leaf1
Kaśka a PL 92 (PlantiCo Hodowla i
Nasiennictwo Ogrodnicze Zielonki
sp. z o.o.)
Root2
Kinga a PL 67 (Krakowska Hodowla i
Nasiennictwo Ogrodnicze POLAN
sp. z o.o.)
Root1
Konika a CZ 1495 (Moravoseed CZ a.s.) Root1
Korai cukor a HU 151508 (ZKI
Zöldségtermesztési Kutató Intézet
Zrt.)
Leaf1
Krista a CZ 256 (Seva-Flora s.r.o.) Leaf1
Kudrnka a CZ 239 (SEMO a.s.) Leaf1
Laica a DE 2549 (Hild Samen GmbH) Leaf1
Lange Oberlaaer a AT 72 (Austrosaat Österreichische
Samenzucht- und Handels-AG)
Root1
Laura a DE 2549 (Hild Samen GmbH) Leaf1
Lenka a PL 1087 (Przedsiębiorstwo
Nasiennictwa Ogrodniczego i
Szkółkarstwa w Ożarowie
Mazowieckim Spółka z o.o.)
Root1
Lisette a DE 2549 (Hild Samen GmbH) Leaf1
Makói hosszú b HU 191126 (Fekete János) Root2
Marunka a CZ 239 (SEMO a.s.) Leaf1
Messis a PL 938 (AdvanSeed ApS) Leaf1
Table 13.2 (continued)
(continued)
13 Petroselinum crispum (Mill.) Nyman (Parsley)
460
Table 13.2 (continued)
Variety name Country maintainers Synonym
Used plant
part (1, 2)
Mooskrause 2
Extra triple
curled 2
Riccio
muschiato 2
Riccio verde
scuro 2
Moskrul 2
Moss Curled 2
Nain frisé
mousse
Nano ricciuto 2
b NL x
b UK x
b FR x
Leaf1
Moskrul 2 = Mooskrause 2 Leaf1
Moss Curled 2 = Mooskrause 2 Leaf1
Nain frisé mousse = Mooskrause 2 Leaf1
Nano ricciuto 2 = Mooskrause 2 Leaf1
Natalka a PL 92 (PlantiCo Hodowla i
Nasiennictwo Ogrodnicze Zielonki
sp. z o.o.)
Leaf1
Nutka a PL 1087 (Przedsiębiorstwo
Nasiennictwa Ogrodniczego i
Szkółkarstwa w Ożarowie
Mazowieckim Spółka z o.o.)
Leaf2
Olomoucká
dlouhá
Ołomuńcka
a CZ x, a SK 250 (Zelseed spol. s
r.o.)
a PL 218 (Przedsiębiorstwo
Nasiennictwa Ogrodniczego i
Szkółkarstwa)
Root1
Oltis a RO 1033 (Staţiunea de Cercetare-
Dezvoltare pentru Legumicultură
Buzău)
Root2
Orbis a CZ 239 (SEMO a.s.) Root1
Orfeo a NL 26 (Enza Zaden Seed
Operations BV)
Leaf1
Ory a RO 1033 (Staţiunea de Cercetare-
Dezvoltare pentru Legumicultură
Buzău)
Leaf1
Osborne a CZ 1495 (Moravoseed CZ a.s.) Root1
Ołomuńcka = Olomoucká dlouhá Root1
Peione a NL 26 (Enza Zaden Seed
Operations BV)
Leaf1
Pesto a PL 356 (Hortag Seed Co.) Leaf2
Petronelia a NL 108 (Rijk Zwaan Zaadteelt en
Zaadhandel BV)
Leaf1
Petruschka a DE 7092 (GHG Saaten GmbH) Leaf1
Plain Leaved
(Sheeps) 2
= Commun 2 Leaf1
Plain or single 2 = Commun 2 Leaf1
(continued)
F. M ar the
461
Variety name Country maintainers Synonym
Used plant
part (1, 2)
Polina a NL 108 (Rijk Zwaan Zaadteelt en
Zaadhandel BV)
Leaf2
Prairie a NL 8 (Bejo Zaden BV) Leaf2
Riccio muschiato
2
= Mooskrause 2 Leaf1
Riccio verde scuro
2
= Mooskrause 2 Leaf1
Rizado verde
oscuro
= Frisé vert foncé Leaf1
Roksana a PL 92 (PlantiCo Hodowla i
Nasiennictwo Ogrodnicze Zielonki
sp. z o.o.)
Root2
Rosette b NL 136 (A.L.Tozer Ltd.) Leaf1
Samba a PL 187 (SPÓJNIA Hodowla i
Nasiennictwo Ogrodnicze sp. z o.o.)
Root1
Simple 2 = Commun 2 Leaf1
Sonata a PL 187 (SPÓJNIA Hodowla i
Nasiennictwo Ogrodnicze sp. z o.o.)
Root1
Starke b DK 56 (Weibulls Horto AB) Leaf1
Starlett a DE 2549 (Hild Samen GmbH) Leaf1
Thujade b NL 65 (Pieterpikzonen BV) Leaf1
Titan b NL 8 (Bejo Zaden BV) Leaf1
Toscano 2 = Commun 2 Leaf1
Troja a CZ 239 (SEMO a.s.) Root2
Verde scuro
d’Italia
= Gigante di Napoli Leaf1
Vistula a PL 67 (Krakowska Hodowla i
Nasiennictwo Ogrodnicze POLAN
sp. z o.o.)
Root2
Walser Petersilie b AT 148 (Arche Noah) Leaf2
Warta a PL 187 (SPÓJNIA Hodowla i
Nasiennictwo Ogrodnicze sp. z o.o.)
Root1
Weg a a NL 26 (Enza Zaden Seed
Operations BV)
Leaf1
Zaharat a RO 1031 (Staţiunea de Cercetare-
Dezvoltare pentru Legumicultură
Bacău)
Leaf1
(1): Variety denomination approved in the form of a code
a: Basic seed
b: Standard seed
Country code: AT– Austria, CZ– Czech Republic, DE– Germany, DK– Denmark, ES– Spain,
FR– France, HR– Croatia, HU– Hungary, IT– Italy, NL– the Netherlands, PL– Poland, RO–
Romania, SK– Slovakia, UK– United Kingdom
Used plant parts
1 (Kraus-Schierhorn 2019)
2 (Blüthner 2019)
Table 13.2 (continued)
13 Petroselinum crispum (Mill.) Nyman (Parsley)
462
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F. M ar the
Handbook of Plant Breeding
JohannesNovak
Wolf-DieterBlüthner Editors
Medicinal,
Aromatic and
Stimulant Plants
Handbook of Plant Breeding
Volume 12
Editor-in-Chief
IstvanRajcan
Department of Plant Agriculture
University of Guelph
Guelph,ON,Canada
JohannVollmann
Department of Crop Sciences
University of Natural Resources and Life Sciences
Vienna,Wien,Austria
The eld of plant breeding covers a broad range of different species and categories
of plants. While there are many techniques and issues that are similar across these
species, there are many more that are unique to each category of species.
The Handbook of Plant Breeding is organized by major crop categories and includes
the most up-to-date molecular techniques being used. It will serve as a resource for
plant breeding laboratories in both the university and industrial setting.
More information about this series at http://www.springer.com/series/7290
Johannes Novak • Wolf-Dieter Blüthner
Editors
Medicinal, Aromatic
and Stimulant Plants
ISSN 2363-8478 ISSN 2363-8486 (electronic)
Handbook of Plant Breeding
ISBN 978-3-030-38791-4 ISBN 978-3-030-38792-1 (eBook)
https://doi.org/10.1007/978-3-030-38792-1
© Springer Nature Switzerland AG 2020
This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microlms or in any other physical way, and transmission or information
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does not imply, even in the absence of a specic statement, that such names are exempt from the relevant
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claims in published maps and institutional afliations.
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The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Editors
Johannes Novak
Institute for Animal Nutrition
and Functional Plant Compounds
University of Veterinary Medicine Vienna
Vienna, Austria
Wolf-Dieter Blüthner
Dr. Junghanns GmbH
Aschersleben, Germany
v
Preface
Medicinal and aromatic plants (MAPs) and stimulant plants include many thousand
plant species with specic physiological effects of their plant secondary compounds
on health, taste of food, and well-being. They can be found in all plant families
worldwide, and their use is not only restricted to humans but also extended more
and more to animal husbandry and plant protection.
The detection and use of these effects by humans date back to ancient times
based on trial and error and are region-specic. However, during history, important
plants and their application were exchanged between cultures and have been incor-
porated into world knowledge.
A major characteristic of MAPs is their richness in species. Whereas less than 5
food plants (sugarcane, rice, wheat, corn, potato) (Joy etal. 1998) save more than
half of the world harvested yield, MAPs comprise between 52,000 species (out of
422,000 owering plant species). In Germany, for example, 1,543 species are
traded, but only 50–100 of these are exclusively sourced from cultivation.
Extrapolating from the gure, only a few hundred species are under cultivation
(Schippmann etal. 2002). With increasing pressure on natural populations, domes-
tication is promoted, and the number of cultivated species will increase. Therefore,
breeding different wild MAP species will increase in the short term.
In food plants, only a few chemical groups with a modest number of substances
are used, namely, carbohydrates, fatty oils, and proteins, while in MAPs, several
hundred thousand different products of the secondary metabolism are of value.
However, with increasing popularity of polyphenols as antioxidants, breeding is
trying to enrich food, vegetables, and fruits with benecial compounds for healthier
and tastier food. So, the borders between food and MAPs are blurred:
• Aromatic plants (herbs and spices) can still be distinguished from “avor-
improved” food by their property of not having nutritional value. They are in a
strict sense not “food” but change the properties of food.
• Stimulant plants, like nicotine, coffee, cocoa, and tea, are also not food but estimated
due to their stimulating effect, often bringing also complex avors into our diet.
• Medicinal plants improve or maintain health but are sometimes used because of
their avor as food (e.g., herbal teas).
vi
Medicinal plants are used (a) as traditional medicine; (b) as phytomedicines,
registered according to pharmaceutical regulatory requirements; and (c) as food
supplements (botanicals), products based on medicinal plants targeting primarily
health maintenance, which are known from traditional medicine to be safe and
effective (but regulated by food law). So, also here, the borders between medicinal
and food use are blurred.
In legislation, however, we need to think in black and white resulting in pharma-
ceutical law and regulations that are dominantly a “negative law” (to say it bold and
simple, “Everything is forbidden that is not allowed.”) and food law and regulations,
a “positive law” (“Everything is allowed that is not forbidden.”), which can become
quite challenging for breeding, if quality criteria– differently dened for medicine
and food– need to be considered.
Only a few species, where seed sales are able to renance breeding investments, are
in intensive and continuous breeding programs of professional breeding companies. As
a result, only a few cultivars are registered and protected, most selections are used in
closed production systems, and maintenance breeding is restricted to the product life
cycle. For most of the MAP species, breeding is project-based, so time- restricted and
as part of a complex optimization of agricultural production. Here, product sales are
renancing optimization investments. For this group, academia, farmer associations
and sometimes even raw material processors, traders, and product producers are practi-
cally breeding in a pragmatic way. This form of breeding is often performed with only
simple breeding techniques like mass selection. This approach, however, can lead in
many cases to a fast, signicant improvement in just a few generations based on the
wide natural variability in the starting materials collected from the wild.
In this book, a few important genera or species are covered in 17 specic chap-
ters. Three general chapters reect the particularities of MAP research and breed-
ing. Two chapters summarize information on over 2,000 MAP species on available
genetic resources, DNA particularities, and pollination biology. Two chapters can
help entering more efciently in more detailed research and breeding work. Another
chapter focuses on peculiarities of chemical analysis of plant secondary compounds
and some approaches to adapt analytics to breeding requirements.
Under the plethora of available literature on MAPs, breeding is severely
neglected. We hope that this compilation contributes to a wider view on MAPs.
Aschersleben, Germany Wolf-DieterBlüthner
Wien, Austria JohannesNovak
References
Joy PP, Thoma J, Mathew S, Skaria BP (1998) Medicinal plants. Kerala Agricultural University.
Schippmann U, Leaman DJ, Cunningham AB (2002) Impact of cultivation and gathering on bio-
diversity: Global trends and issues. In: FAO (eds) Biodiversity and the ecosystem approach in
agriculture, forestry and sheries. FAO, Rome, pp142–167.
Preface
vii
Contents
1 Genetic Resources of Medicinal and Aromatic Plants . . . . . . . . . . . . . 1
Ulrike Lohwasser and Stephan Weise
2 Analysis of Secondary Metabolites in Breeding Research
and Plant Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Hartwig Schulz
3 Cannabis sativa L. –Cannabis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Paweł Rodziewicz and Oliver Kayser
4 Coriandrum sativum L. – Coriander . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
Axel Diederichsen, Sabine Banniza, Cheryl Armstrong-Cho, and
Travis Sander
5 Duboisia sp. – Corkwood Tree . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
Julia Sparke
6 Lavandula angustifolia Mill. and Lavandula x intermedia
Emeric ex Loisel: Lavender and Lavandin . . . . . . . . . . . . . . . . . . . . . . 303
Philippe Gallotte, Guillaume Fremondière, Philippe Gallois,
Jean- Pierre Bouverat Bernier, Anne Buchwalder, Alan Walton,
Josephine Piasentin, and Berline Fopa-Fomeju
7 Matricaria recutita L.: True Chamomile . . . . . . . . . . . . . . . . . . . . . . . . 313
Sebastian Albrecht and Lars-Gernot Otto
8 Nasturtium officinale R.Br.: Watercress . . . . . . . . . . . . . . . . . . . . . . . . 333
Wolf-Dieter Blüthner
9 Nicotiana tabacum L.: Tobacco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Ramsey S. Lewis
10 Ocimum basilicum L. (Basil). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Nativ Dudai, Nadav Nitzan, and Itay Gonda
viii
11 Origanum majorana L. (Marjoram) . . . . . . . . . . . . . . . . . . . . . . . . . . . 407
Brigitte Lukas and Johannes Novak
12 Origanum vulgare L. and Origanum onites L. (Oregano) . . . . . . . . . . 419
Brigitte Lukas and Johannes Novak
13 Petroselinum crispum (Mill.) Nyman (Parsley) . . . . . . . . . . . . . . . . . . . 435
Frank Marthe
14 Rosa x damascena Mill. (Rose) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 467
Krasimir Rusanov, Natasha Kovacheva, Ana Dobreva, and Ivan
Atanassov
15 Rosmarinus officinalis L.: Rosemary . . . . . . . . . . . . . . . . . . . . . . . . . . . 501
Merita Hammer and Wolfram Junghanns
16 Salvia officinalis L. and Salvia fruticosa Mill.: Dalmatian
and Three- Lobed Sage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 523
Corinna Schmiderer and Johannes Novak
17 Salvia sclarea L.: Clary Sage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539
Berline Fopa-Fomeju, Philippe Gallotte, Philippe Gallois, Guillaume
Fremondière, Jean-Pierre Bouverat Bernier, and Anne Buchwalder
18 Thymus vulgaris L.: Thyme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547
José F. Vouillamoz and Bastien Christ
19 Valeriana officinalis L. s.l.: Valerian . . . . . . . . . . . . . . . . . . . . . . . . . . . 559
Michael Penzkofer and Heidi Heuberger
20 Zingiber officinale Roscoe: Ginger . . . . . . . . . . . . . . . . . . . . . . . . . . . . 605
Meenakshi Kumari, Manoj Kumar, and S. S. Solankey
21 DNA Content (C-Values), Chromosome Numbers,
and Mating System of Medicinal, Aromatic,
and Stimulant Plants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623
Wolf-Dieter Blüthner
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 625
Contents
