Digitalis thapsi

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Digitalis thapsi
Digitalis thapsi 01.JPG
Digitalis thapsi in situ at Villavieja de Yeltes, Salamanca, Spain
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Lamiales
Family: Plantaginaceae
Genus: Digitalis
Species:
D. thapsi
Binomial name
Digitalis thapsi
Synonyms

Homotypic:

  • Digitalis purpurea proles thapsi((L.) Font Quer in Bol. Farm. Militar 3: 128. 1925.)
  • Digitalis purpurea subsp. thapsi((L.) Font Quer, Pl. Medic. 619. 1962, comb. inval.)
  • Digitalis purpurea subsp. thapsi((L.) Rivas Goday in Farmacogn. 5: 144. 1946.)

Heterotypic:

  • Digitalis thapsi f. albiflora(C.Vicioso, in sched., nom. nud. )
  • Digitalis thapsi f. albiflora(C.Vicioso ex R.Fern. in Mem. Soc. Brot. 6: 50. 1950.)

Digitalis thapsi, which has been called mullein foxglove in the US, is a flowering plant in the genus Digitalis that is endemic to the Iberian Peninsula, where it occurs in eastern Portugal and central and western Spain. It is of commercial importance as an ornamental plant. Hybrids with D. purpurea have proved successful and are fertile.

Contents

Names

A Missouri Botanical Garden website calls it mullein foxglove. [2] Local vernacular names recorded for this species are abeloura, abeloura-amarelada, aboleira, aveloeira, dedaleira, luvas de Santa María, pegajo and rabo de raposa in Portuguese, whereas in Spanish the most common name is dedalera, followed by viloria, viluria, giloria, dedales and mataperla, but it has also been called abiloria, abiluria, abortones, bacera, beleño, biloria, biluria, campanilla, cascante, cascaor, chupadera, chupamieles, chupera, cohete, cohetera, cohetes, deales, dedales de niño, dedales purpúreos, dediles, digital, digitalis, emborrachacabras, estallones, goldaperra, guadalperra, gualdaperra, guardaperros, hueltaperra, mata de lagartija, mataperros, rabera, ravera, raéra, restalladera, restralleti, restrallos, sabia, tuercecuellos and vueltaperra. [3] [4]

Taxonomy

It was first described in the modern taxonomic system by Carl Linnaeus in 1763. [1]

The genus Digitalis was formerly assigned to the family Scrophulariaceae, [5] but it is now considered to belong within the Plantaginaceae. [6]

Hybrids

Joseph Gottlieb Kölreuter, a German botanist, observed that D. thapsi had many characteristics of D. purpurea after four to five generations of cultivation, and that the former became indistinguishable from the latter, a report that was considered "probably" trustworthy by Charles Darwin. D. thapsi and D. purpurea hybrids are generally fertile. [7]

In many areas of Spain and Portugal populations of D. thapsi and D. purpurea meet, and introgression is common. These have been known as D. purpurea nothosubsp. carpetana and D. minor in error in two older Portuguese works (1906, 1913), but this hybrid is now generally known as D. × coutinhoi. [8]

Description

Growing in its native Spain Digitalis thapsi 20100522 1.jpg
Growing in its native Spain

Habitus

Digitalis thapsi is a perennial plant. [8] [9] Its yellow-green leaves are ovate to oblong in shape and decurrent, with winged bases. [10] The flowers, stems and leaves are covered with a dense indumentum of tiny hairs, giving them a soft appearance. [8] [10] These yellow-green hairs (trichomes) are exclusively glandular. The entire plant is very sticky (glutinose). It is caespitose, meaning it grows with its large leaves tightly arranged into one, or a number, of rosettes growing at or just above ground level. In older plants the base becomes woody and highly branched at ground level, so that a single plant has multiple rosettes. The green, round to slightly angular stems grow to (30-)50 to 80(-100)cm in height before they start to flower. The stems are similarly entirely covered in glandular hairs; these are present in two forms: short and subsessile, or much longer, 0.4 to 0.6mm. [8]

The chromosome count is 2n=56. [8]

Leaves

The lower basal leaf blades are 7.5 to 13 cm (exceptionally 19 cm) in length, 2.5 to 5 cm in width, more or less flat but sometimes undulated along the margins, and oblong to elliptic in shape, with a sharp to somewhat sharply tipped apex. The bases of the leaves taper gradually into the petioles. The texture is soft, not leathery. The underside is rugose-reticular in texture, coloured greenish-yellow and very densely packed with glandular hairs. The leaves in the middle of the rosette are clearly decurrent. The margins are denticulate or somewhat so, rarely subentire. [8]

Flowers

The pink, rarely white, [8] flowers are arranged in an inflorescence in the form of a raceme, [11] 15 to 35 cm in length with 15 to 40 individual flowers. The inflorescence has glandular hairs along its shaft, is not stiff, and is secundiflorous, meaning the flowers follow each other in succession. The flowers have a 1 to 2 cm long pedicel which is curved at the base, and shorter to equal in length to the subtending bracts. The flowers hang somewhat, and are separated by internodes of 8-15mm. The bracts are 12 to 20mm in length and 2 to 3mm broad, glandular and lanceolate in shape. [8]

Chemistry

According to Dutch botanist Herman Boerhaave, writing at the turn of the 17th century, Digitalis species are highly poisonous if directly ingested. [12] All parts are poisonous. [11] [13] In studies, the restriction of calcium ions resulted in cardenolide accumulation in D. thapsi. [14] Calcium quantity affects the redox chemical reactions in the cells. Without calcium, changes in antioxidant function were observed and catalase activity was slow. [15] In another study, lack of calcium retarded growth and promoted digoxin formation. Manganese sulfate and lithium chloride also increased the digoxin concentration, but did not affect growth. [16] Calcium oxalate crystals have also been isolated from D. thapsi. [10] [17] [18]

Similar species

In 1841 Pierre Edmond Boissier considered it the most similar to D. mariana, which he was then describing as a new species, having collected from the same region. [19] In the dichotomous key in the Flora Iberica , it is keyed out to D. mariana, D. minor and D. purpurea. D. thapsi is distinguished from these three other similar species by being the only very sticky species, being completely covered in relatively long, yellowish, glandular hairs. It is also the only species in which the leaves in the middle of the rosette are clearly decurrent. D. minor, an endemic of the Balearic Islands, is not sympatric with D. thapsi. [8]

Distribution

Digitalis thapsi is an endemic species of the Iberian Peninsula, [4] [6] occurring in both Portugal and Spain. It grows in the mountains and rocky plains of the central and central-western parts of the Iberian Peninsula, especially in the Sistema Central and its extensions between the rivers of the Douro and Tagus. [8]

It is widespread in Spain, [5] occurring in the western and central parts of the country. [6] [20] [21] It does not occur on the Balearic Islands. [4] It occurs in the provinces of Ávila, Badajoz, Cáceres, Córdoba, Guadalajara, Madrid, Salamanca, Segovia, Soria, Toledo and Zamora. It has incidentally been found in the province of Burgos. [8] In Andalucía it is only found in the north of the region, in the western Sierra Morena, where it is uncommon; [22] it has been collected more often in the mountains of central Spain. [3] [8]

In Portugal it primarily occurs in the northern and central interior, [6] [21] more precisely in the districts of Bragança, Vila Real, Guarda, Viseu, Castelo Branco, Portalegre and Évora. With less frequency, it has also been found in the Porto, Aveiro, Coimbra, Santarém, Beja and Faro Districts. [8] [23]

Ecology

Digitalis thapsi grows in acidic soils. [22] It grows on rocky ground [8] [22] and on slopes. It prefers sunny, open and disturbed places. It generally grows on soils derived from granite, but also sometimes on quartzite and schist. It grows at (rarely 100-) 500–1500 metres in altitude. [8]

The specific phytosociological suballiance and alliance wherein it occurs is called 'Rumici indurati-Dianthion lusitani', which is in the order 'Phagnalo saxatilis-Rumicetalia indurati', in the class 'Phagnalo saxatilis-Rumicetea indurati'. It is a characteristic species of this phytocoenosis. In this habitat it occurs together with these following characteristic species: Antirrhinum graniticum subsp. graniticum, Antirrhinum rupestris , Arrhenatherum fernandesii , Biscutella bilbilitana , Centaurea monticola subsp. citricolor, Centaurea pinnata , Coincya leptocarpa , Coincya longirostra , Coincya pseudoerucastrum subsp. pseudoerucastrum, Coincya rupestris , Conopodium bunioides subsp. aranii, Conopodium majus subsp. marizianum, Crepis oporinoides , Dianthus crassipes subsp. crassipes, Dianthus crassipes subsp. sereneanus, Dianthus lusitanus , Digitalis mariana (both subspecies), Digitalis purpurea subsp. toletana, Erodium mouretti , Erysimum linifolium subsp. lagascae, Festuca duriotagana , Jasione sessiliflora subsp. tomentosa, Scrophularia oxyrhincha , Scrophularia sublyrata , Sedum hirsutum subsp. baeticum, Silene marizii , Silene × montistellensis and Verbascum rotundifolium subsp. castellanum. [24]

In its native lands it flowers from June to July. [8] In cultivation it flowers in late spring [2] [11] to early summer. [2] A study of the fruiting activity in D. thapsi showed that production, size and number of seeds were interrelated. Analysis has shown that seed weight is not related to the length of the cotyledon, and that the length of the fifth leaf can serve as a useful factor in determining the genetic variability among specimens. [5]

Uses

Agriculture

It is known that livestock avoid eating this plant, but an exception are goats, which will eat the leaves and flowers in times of shortage. Nonetheless, the plant is poisonous for them, causing stupor and paralysis -this was apparently once not an uncommon occurrence in Spain, and has spawned a number of idioms. [25]

Cultivation

The species is used as a perennial, ornamental plant. [9] It is low-growing, so is used in the border. It grows well in half-shaded to sun-exposed areas in temperate regions. [2] [11] It prefers organically rich, reasonably well-drained, acidic soils. In Missouri it is said to require constantly moist soil. [2] In British gardens moderate watering is required; the plant has an average drought tolerance, [11] and is adaptable to dry shady areas. [26] It is tolerant of deer. Many gardeners remove the spent flower spikes soon after bloom, not allowing it to go to seed. [2]

Besides normal cultivation, D. thapsi has been propagated using explant culture, a technique employed by isolating and harvesting meristem cells from pieces of tissue. Auxins NAA, 2,4-D and IAA alone or combined with BA produced a callus. NAA caused root formation and BA shoot formation. NAA and BA combined induced organ generation more effectively. Plantlets obtained this way had a survival rate of 70%. [27]

A cultivar called 'Spanish Peaks' with raspberry-rose-coloured flowers and a compact habitus has been available in the US. [2]

Traditional

In Spain the traditional uses have practically been abandoned. Recorded traditional uses are often identical to that those of D. purpurea; when questioned many informants in Salamanca believed that these were in fact the same species. It is generally thought that the difference in flower colour is due to some characteristic of the soil. Local people are aware it is poisonous. It was once locally used in herbalism as a cardiac tonic in many places. Other folk medicinal uses are local to specific cities or villages: in the hills near Salamanca the leaves were steeped in water to use for a sore throat or a decoction used for infected wounds, in a town in Galicia the flowers in water were used to combat inflammations, in the provinces of Zamora and Salamanca boils are said to heal after applying a poultice of the burnt basal leaves. [25]

Conservation

It is not considered rare in a national or international perspective, [3] but it has been classified as locally 'endangered' in the Regional Red List of Andalucía of 2005. [3] [22] The reason for this was that the different collection localities and populations in this region were fragmented and very small. [22]

Related Research Articles

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<i>Digitalis</i> Genus of flowering plants in the family Plantaginaceae

Digitalis is a genus of about 20 species of herbaceous perennial plants, shrubs, and biennials, commonly called foxgloves.

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<span class="mw-page-title-main">Wildflower</span> Flower that grows in the wild, not intentionally planted

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<i>Digitalis purpurea</i> Toxic flowering plant in the family Plantaginaceae

Digitalis purpurea, the foxglove or common foxglove, is a toxic species of flowering plant in the plantain family Plantaginaceae, native to, and also widespread throughout most of temperate Europe. It has also naturalized in parts of North America, as well as some other temperate regions. The plant is a popular garden subject, with many cultivars available. It is the original source of the medicine digoxin, a medicine for the human heart. This biennial plant grows as a rosette of leaves in the first year after sowing, before flowering and then dying in the second year. It generally produces enough seeds, however, so that new plants will continue to grow in a garden setting.

<i>Pilosella officinarum</i> Species of flowering plant in the daisy family Asteraceae

Pilosella officinarum, known as mouse-ear hawkweed, is a yellow-flowered species of flowering plant in the daisy family Asteraceae, native to Europe and northern Asia. It produces single, lemon-coloured inflorescences. Like most hawkweed species, it is highly variable and is a member of a species complex of several dozens of subspecies and hundreds of varieties and forms. It is an allelopathic plant.

<i>Dianthus armeria</i> Species of flowering plant

Dianthus armeria, the Deptford pink or grass pink, is a species of Dianthus ("pink") native to most of Europe, from Portugal north to southern Scotland and southern Finland, and east to Ukraine and the Caucasus. It is naturalised in North America.

<i>Digitalis lanata</i> Species of flowering plant in the plantain family Plantaginaceae

Digitalis lanata, vernacularly often called woolly foxglove or Grecian foxglove, is a species of foxglove, a flowering plant in the plantain family Plantaginaceae. It gets its name due to the woolly indumentum of the leaves. D. lanata, like other foxglove species, is toxic in all parts of the plant. Symptoms of digitalis poisoning include nausea, vomiting, severe headache, dilated pupils, problems with eyesight, and convulsions at the worst level of toxicity. The plant is also harmful to other animals.

<i>Digitalis grandiflora</i> Species of foxglove

Digitalis grandiflora, the yellow foxglove, big-flowered foxglove, or large yellow foxglove, is a species of flowering plant in the genus Digitalis, family Plantaginaceae. It is native to southern Europe and Asia. In mountains it grows on warm, bushy slopes or areas left after logging. The Latin specific epithet grandiflora means “large flowered”.

<i>Olearia phlogopappa</i> Species of flowering plant

Olearia phlogopappa commonly known as the dusty daisy-bush or alpine daisy-bush is a species of flowering plant in the family Asteraceae that is commonly found in eastern New South Wales, Victoria and Tasmania. It is a small shrub with greyish-green foliage, daisy-like flowers in white, pink or mauve that can be seen from spring to late summer.

<i>Pinguicula orchidioides</i> Species of carnivorous plant

Pinguicula orchidioides is a perennial rosette-forming insectivorous herb native to Mexico and Guatemala. A species of butterwort, it forms summer rosettes of flat, succulent leaves up to 5 centimeters (2 in) long, which are covered in mucilaginous (sticky) glands that attract, trap, and digest arthropod prey. Nutrients derived from the prey are used to supplement the nutrient-poor substrate that the plant grows in. Uniquely among Pinguicula species from the Americas, p. orchidioides produces gemma-like basal buds which elongate into stolons and serve as a means of asexual reproduction. In the winter the plant forms a non-carnivorous rosette of small, fleshy leaves that conserves energy while food and moisture supplies are low. Single purple flowers appear between July and September on upright stalks up to 22 centimeters long.

<i>Digitalis <span style="font-style:normal;">×</span> fulva</i> Hybrid species of flowering plant

Digitalis × fulva, the strawberry foxglove, is a hybrid species of flowering plant within the family Plantaginaceae. It is a naturally occurring fertile hybrid between the species Digitalis grandiflora and Digitalis purpurea. The species is widely marketed in the UK under the common name strawberry foxglove or its taxonomic synonym Digitalis × mertonensis. The species has been used to produce various cultivars and has gained the Royal Horticultural Society's Award of Garden Merit.

<i>Digitalis ciliata</i> Species of plant

Digitalis ciliata, commonly called hairy foxglove is a member of the genus Digitalis. It has thimble-shaped, yellow to cream colored flowers produced on perennial plants with evergreen foliage. It is native to the Caucasus and is grown as an ornamental in other parts of the world. The species name is derived from the fine hairs that cover the plants stems and flowers.

<i>Digitalis obscura</i> Species of plant

Digitalis obscura, commonly called willow-leaved foxglove or dusty foxglove or spanish rusty foxglove, is a flowering plant native to regions in Spain and Morocco. It is also grown as an ornamental flower. This foxglove is a woody perennial plant belonging to the family Plantaginaceae. Along with the other foxgloves it used to be placed in the figwort family, Scrophulariaceae; however, recent genetic research has moved the genus Digitalis to a larger family. It is similar to many of the foxglove species in its high toxicity and medicinal use as a source for the heart-regulating drug digoxin. Its strikingly distinctive amber- to copper-coloured flowers give the species its name and help distinguish it from other members of the genus.

<i>Goodenia heterophylla</i> Species of flowering plant

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<i>Petrorhagia saxifraga</i> Species of flowering plant

Petrorhagia saxifraga, known as tunic flower or coat flower, is a small, herbaceous flowering plant in the family Caryophyllaceae. It is native to parts of Europe and introduced to the United States and Canada, Great Britain, and Sweden. Petrorhagia saxifraga is also known as tunic saxifrage, pink saxifrage, or just pink.

<i>Androsace vitaliana</i> Species of flowering plant

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<i>Digitalis mariana</i> Flowering plant in the family Plantaginaceae

Digitalis mariana is a flowering plant species in the family Plantaginaceae. It is a perennial foxglove with evergreen foliage and rose-red coloured flowers produced in summer. It is native to Portugal and Spain.

<i>Digitalis minor</i> Species of plant

Digitalis minor is a species of flowering plant in family Plantaginaceae, which has been called dwarf Spanish foxglove. It is a biennial or short-lived perennial species of foxglove which is endemic to the Balearic islands with large, pendulous, pink or purple flowers. Closely related to the common purple foxglove, it is best distinguished by its small fruits. It is one of the only foxgloves to grow in calciferous, alkaline soils.

<i>Digitalis nervosa</i> Species of flowering plant

Digitalis nervosa is a species of flowering plant in family Plantaginaceae. It is native to the Caucasus down to north western and northern Iran.

References

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  2. 1 2 3 4 5 6 7 "Digitalis thapsi 'Spanish Peaks'". Plant-finder. Missouri Botanical Garden . Retrieved 31 August 2012.
  3. 1 2 3 4 "Búsquedas: Digitalis thapsi". ANTHOS (in Spanish). Fundación Biodiversidad, Ministerio de Agricultura, Alimentación y Medio Ambiente, & Real Jardín Botánico. 2 March 2012. Retrieved 23 November 2020.
  4. 1 2 3 Marhold, Karol (2011). Greuter, Werner; von Raab-Straube, E. (eds.). "Details for: Digitalis thapsi". Euro+Med Plantbase. Botanic Garden and Botanical Museum Berlin-Dahlem . Retrieved 23 November 2020.
  5. 1 2 3 Cabello, M. L.; Ruiz, T.; Devesa, J. A. (2001). "Carpology, germination and seedling performance in Digitalis thapsi L. (Scrophulariaceae)". Botanical Journal of Scotland. 53 (2): 135–154. doi:10.1080/03746600108685019. S2CID   84615329.
  6. 1 2 3 4 "Digitalis thapsi L." Plants of the World Online . Royal Botanic Gardens, Kew. 2017. Retrieved 23 November 2020.
  7. ↑ Stauffer, R. C. (1987). Charles Darwin's Natural selection : being the second part of his big species book written from 1856 to 1858 (1, paperback ed.). Cambridge: Cambridge University Press. p.  127. ISBN   978-0-521-34807-2.
  8. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Benedí i Gonzalez, Carles; Hinz Alcaraz, P.-A. (15 January 2009). "17. Digitalis" (PDF). In Benedí i Gonzalez, Carles; Rico Hernández, Enrique; Güemes Heras, Jaime; Herrero Nieto, Alberto (eds.). Flora Ibérica, Vol. XIII (in Spanish). Madrid: Real Jardín Botánico. pp. 342–343, 351–353, 357. ISBN   9788400087470.
  9. 1 2 Kole, Chittaranjan (1 September 2011). Wild Crop Relatives: Genomic and Breeding Resources: Plantation and Ornamental Crops. Springer Science & Business Media. p. 78. ISBN   978-3-642-21201-7.
  10. 1 2 3 Kar, A. (2003). Pharmacognosy And Pharmacobiotechnology. New Delhi: New Age International Ltd. p. 173. ISBN   978-81-224-1501-8.
  11. 1 2 3 4 5 "Digitalis thapsi". Plant Database. Archived from the original on 6 January 2014. Retrieved 2 October 2012.
  12. ↑ Encyclopædia Britannica. Vol. 4 (6th ed.). Edinburgh: Archibald Constable and Company. 1823.
  13. ↑ Edinger, Philip; Sanchez, Janet H. (2002). Annuals and Perennials . Sunset Books. p.  118.
  14. ↑ Cacho, M. M.; et al. (1995). "Calcium restriction induces cardenolide accumulation in cell suspension cultures of Digitalis thapsi L". Plant Cell Reports. 14 (12): 786–789. doi:10.1007/bf00232923. ISSN   0721-7714. PMID   24186713. S2CID   8040006.
  15. ↑ Paranhos, A.; Fernandez-Tarrago, J.; Corchete, P. (1 January 1999). "Relationship between active oxygen species and cardenolide production in cell cultures of Digitalis thapsi: effect of calcium restriction". New Phytologist. 141 (1): 51–60. doi:10.1046/j.1469-8137.1999.00317.x.
  16. ↑ Corchete, M. P.; et al. (1 October 1991). "Effect of calcium, manganese and lithium on growth and cardenolide content in cell suspension cultures of Digitalis thapsi L.". Plant Cell Reports. 10 (8): 394–396. doi:10.1007/BF00232609. ISSN   1432-203X. PMID   24221731. S2CID   21383174.
  17. ↑ Kokate, C. K.; Gokhale, S. B.; Purohit, A. P. (2009). Pharmacognosy. Nirali Prakashan. pp. 39–40. ISBN   978-81-963961-5-2 . Retrieved 31 August 2012.
  18. ↑ Khandewal, K. R. (2008). Practical Pharmacognosy. Pune: Nirali Prakashan. p. 59. ISBN   978-81-85790-30-5.
  19. ↑ Boissier, Pierre Edmond (1841). Voyage botanique dans le midi de l'Espagne (in French and Latin). Vol. II. Paris: Gide et Cie. p. 466.
  20. ↑ "Digitalis thapsi". Germplasm Resources Information Network . Agricultural Research Service, United States Department of Agriculture . Retrieved 21 January 2018.
  21. 1 2 Hassler, M. (November 2018). "Synonymic Checklists of the Vascular Plants of the World, Species Details : Digitalis thapsi L." World Plants. Digital resource at www.catalogueoflife.org/col. ISSN   2405-8858 . Retrieved 21 November 2020.
  22. 1 2 3 4 5 Cabezudo, B.; Talavera, S.; Blanca, G.; Salazar, C.; Cueto, M. J.; Valdés, B.; Hernández Bermejo, J. E.; Herrera, C.; Rodríguez Hiraldo, C.; Navas, D. (2005). Lista roja de la flora vascular de Andalucía (PDF) (in Spanish). Sevilla: Consejería de Medio Ambiente, Junta de Andalucía. p. 32. ISBN   84-96329-62-3.
  23. ↑ "Digitalis thapsi L." Flora-On . Retrieved 2 August 2022.
  24. ↑ Menéndez Valderrey, Juan Luis (27 February 2017). "Num. 622: Digitalis thapsi L." Asturnatura.com (in Spanish). AsturnaturaDB. ISSN   1887-5068 . Retrieved 24 November 2020.
  25. 1 2 González, José Antonio; Vallejo, José Ramón; Amich, Francisco (May 2018). "Digitalis thapsi L.". In Pardo de Santayana, Manuel; Morales, Ramón; Tardío, Javier; Molina, Maria (eds.). Inventario Español de los Conocimientos Tradicionales Relativos a la Biodiversidad (in Spanish). Vol. II (1 ed.). Ministerio de Agricultura y Pesca, Alimentación y Medio Ambiente (MAPAMA). pp. 312–313.
  26. ↑ Cullen, James; Knees, Sabina G.; Cubey, H. Suzanne; Shaw, J. M. H. (11 August 2011). The European Garden Flora Flowering Plants: A Manual for the Identification of Plants Cultivated in Europe, Both Out-of-Doors and Under Glass. Cambridge University Press. p. 211. ISBN   978-0-521-76164-2.
  27. ↑ Cacho, Margarita; Morán, Margarita; Herrera, María Teresa; Fernández-Tárrago, Jorge (May 1991). "Morphogenesis in leaf, hypocotyl and root explants of Digitalis thapsi L. cultured in vitro". Plant Cell, Tissue and Organ Culture. 25 (2): 117–23. doi:10.1007/BF00042182. S2CID   41305774.
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