Hypericum perforatum

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[1] [2]

Propagation

Germination

media germination temperature °C note citation
filter paper 52.33 (16.00-75.33) % 18°C light; germination treatments [3]
filter paper 8.78 (1.00-31.33) % 18°C dark; germination treatments [3]

Germination pretreatment efficacy is variable between different accessions of perforatum.[4]

It was determined that the calcium ions present in tap water significantly slow the germination of perforatum. At 7 days, the germination rate was only 45% when hydrated with tap water compared to 92% when distilled water is used. The effect was minimized over time but had not completely disappeared by the end of the study period at 9 days. The effect was distinct at even 1% tap water diluted in distilled water and for temporary immersion in tap water followed by distilled.[5] This effect might be caused by phosphate and iron lockout.[6] However, later studies could not replicate the effect of calcium on germination.[7]

Seeds from plants grown with at least a 17-hour photoperiod had a 96% germination rate compared to 46% with only a 15-hour photoperiod.[8]

Mean germination time was reduced from 28 days in controls to less than 19 days with either 150 mM GA3 or 150 mM KNO3 pretreatment. Similarly, germination fraction increased from 28% to 70% and 71%, respectively.[9]

Positive photoblastic.[10][3]

Seeds can be direct-sown in the field. Combining the tiny seeds with sifted dry peat (1:10) can assist with broadcasting. Germination occurs after 21-25 days at 18-25°C.[10]

[3]

Vegetative

In-Vitro

basal media supplements source target note reference
           

Cultivation

Planting density (m-2) inter-row space (cm) intra-row space (cm) note reference
6.6 50 30 infraspecific variation [11]
10 40 25 fertilization [12]

[10]

Harvest

Perforatum should be harvested when no more than 20% of the flowers have opened to maximize hypericin content.[13]

Because the variation in active constituents varies by more than an order of magnitude between aerial parts of the plants, the harvest ratio of these parts should be controlled to ensure a standardized product.[13] One method is to harvest everything above the “flowering horizon” established by the plane of the lowest flowers. This can vary from 12 to 34cm above the ground according to harvest time, accession, and growing location.[14]

Hypericin, hyperforin, and pseudohypericin content are not affected by drying temperature within the range of 40-80°C. However, the higher temperatures do reduce the flavone glycoside content considerably.[14]

Yield

product source yield per season (kg/ha) note reference
dry biomass seed 100-400 review [10]
dry biomass aerial part 1500-5200 review [10]
dry biomass aerial part 3800-5800 harvest period [13]
product source yield per plant note reference
fresh biomass top 30cm part 3.2 (1.73-5.44) [15]  

Plants may not achieve flowering during the first year of cultivation.[15]

[10] [13] [11] [12]

Soilless

[16]

Soil

soil type pH C-content % precipitation temperature (°C) altitude (m) note reference
               

Seedlings can be transplanted to the field when they are 25cm tall.[12]

[13] [11] [12] [15] [17]

Fertilization

type rate time note reference
urea 0/150/250 kg N/ha pre-transplant fertilization [12]
super phosphate 0/100/200 kg P/ha pre-transplant fertilization [12]

Fertilization with nitrogen and phosphorus increases the dry herbage yield and the hypericin content of perforatum. However, high fertilization also increased the accumulation of cadmium. It is unclear if this is merely a side effect of the natural contamination of commercial mineral fertilizers with cadmium.[12]

[13] [11] [12] [15] [17]

Temperature

Lighting

fixture type photoperiod illumination note reference
         

Supplemental lighting can increase the number of flowers, seeds per flower, seeds per capsule, capsules per plant, flowers per branch, and germination rate. Extending the daylight with supplemental lighting to 17 hours was sufficient to induce these changes, though the highest quality seeds were produced with a 19-hour photoperiod. Alternatively, one hour of night interruption with 30 umol/m2/s fluorescent lamps produced similar, but not identical, changes.[8]

Pests

Rhizoctonia solani infections can wipe out an entire crop.[13]

Powdery mildew attacks fields of perforatum with varying susceptibility among lines.[14]

[13] [12] [14]

Ecology

Morphology

character measurement unit notes reference
height 83.19 (67.60-92.36) cm 1st year; infraspecific variability [15]
inflorescence length 26.13 (16.90-30.50) cm    
inflorescence width 12.52 (9.2-15.5) cm    
leaf length (6th node) 25.43 (18.4-31.30) mm    
leaf width (6th node) 9.26 (6.70-12.10) mm    
petal length 11.98 (9.90-13.60) mm    
petal width 6.17 (5.10-7.00) mm    
sepal length 5.83 (4.40-6.90) mm    
sepal width 1.45 (1.10-1.90) mm    
height 59.98 (41.53-70.55) cm 2nd year; infraspecific variability [15]
inflorescence length 24.21 (21.73-29.07) cm 2nd year; infraspecific variability [15]
inflorescence width 11.36 (8.67-12.93) cm 2nd year; infraspecific variability [15]
leaf length (6th node) 23.60 (18.60-29.10) mm 2nd year; infraspecific variability [15]
leaf width (6th node) 9.49 (7.40-12.20) mm 2nd year; infraspecific variability [15]
petal length 13.35 (11.07-15.80) mm 2nd year; infraspecific variability [15]
petal width 6.17 (5.53-6.87) mm 2nd year; infraspecific variability [15]
sepal length 5.82 (5.33-6.27) mm 2nd year; infraspecific variability [15]
sepal width 1.86 (1.27-2.30) mm 2nd year; infraspecific variability [15]

One out of thirty-one accessions of wild perforatum are hexaploid. The rest are tetraploid. No distinct phenotype was observable from the difference.[18] The highest hypericin content is found in diploids.[2]

Aerial parts of perforatum produce microscopic (89-328 um) black nodules rich in alkaloids and tannins.[19] The presence or absence of these nodules on specific plant parts can be used to group perforatum into morphotypes.[17]

[16] [11] [17] [15] [14] [15]

Roots

Stem

Leaves

Inflorescence

Flowers open at sunrise and last 12-36 hours.[14]

Seeds

Seed pod maturation takes about three weeks.[14]

Phytochemistry

compound source concentration (mg/g dry weight) note citation
hypericin flowers 1.94-2.50 harvest period [13]
hypericin leaves 0.6-0.9 harvest period [13]
hypericin stems 0.07-0.08 harvest period [13]
hypericin aerial parts 0.06-5.2 infraspecific variation [14]
total extract aerial parts 162-358 infraspecific variation [14]
hypericin flowering tops 0.60 (0.23-1.24) infraspecific variation [17]
rutin flowering tops 3.17-13.43 infraspecific variation [17]
hyperoside flowering tops 7.72-31.13 infraspecific variation [17]

[13] [11] [17]

Infraspecific Variation

Hypericin content varies by almost two orders of magnitude between breeding lines of perforatum.[14]

[11] [17] [15] [14]

Biosynthesis

The presence and abundance of black nodules on perforatum are correlated with hypericin content.[17]

Distribution

Hypericin content of the flowers, leaves, and stems is present in the approximate ratio of 30:10:1.[13]

Timecourse

Improvement

trait improvement status reference
     

Current breeding programs are focused on increasing hypericin/hyperforin content and disease resistance.[2]

Identification

variety description reference
Topas(z) Polish standard cultivar [12][14][15]
Zolotodolinskaya Russian [17]

[17] [15] [14]

Inheritance

Methods

type note reference
     

Perforatum can be self-pollinated but will only produce a small seed set.[14]

Faculative, Pseudogamous, apomictic.[2]

[14]

History & Society

Work Log

11 Feb 2023

Seedlings of perforatum are slow growing. I decided to start some seeds today to get a head start. This had nothing to do with my impatience to start the 2023 growing season. Definitely not.

Started 24 seeds of PI664859. Placed directly on the surface of 6-cell plastic flats of PPPM1 then watered from above. I chose this number semi-randomly. These are old seeds with an unknown germination rate.

There are no grazing animals in the immediate area where these plants will go.

04 Apr 2022

First germination:

25 Mar 2022

12 seeds started. Disinfected in the usual manner: 70% EtOH 1 minute + 0.6% hypochlorite for 4 minutes. Sowed onto filter paper in a Petri dish in the 30°C incubator.

09 Nov 2020

A flower from a previous attempt at growing perforatum.

Bibliography

  1. Barnes, Joanne and Arnason, John T and Roufogalis, Basil D, St John's Wort (Hypericum Perforatum L.): Botanical, Chemical, Pharmacological and Clinical Advances, Journal of Pharmacy and Pharmacology, vol. 71, no. 1, pp. 1--3, December 2018. doi: 10.1111/jphp.13053.
  2. Poutaraud, A. and Girardin, P., Improvement of Medicinal Plant Quality: A Hypericum Perforatum Literature Review as an Example, Plant Genetic Resources, vol. 3, no. 2, pp. 178--189, August 2005. doi: 10.1079/PGR200567.
    Numerous factors influence the chemical quality of medicinal plants from crop establishment to extraction of raw material. The most important ones are described using the example of Hypericum perforatum. Optimization of these factors contributes to the objective of producing a high-quality drug, and a method consisting of three scientific approaches (technological, agronomical, plant breeding) is presented. All data concerning the plant (biology, physiology and environmental impacts) and the active components and by-products (pathway, localization and stability) are useful to adapt and to develop management sequences. Although plant breeding appears to be the principal way of improvement, and gives good results in terms of resistance to pathogens, active component content and yield; the agronomical and the technological approaches are also very important. The technological approach after harvesting is especially important to avoid degradation of the active components and to induce, in some cases, the transformation of by-products to those molecules sought. This integrated method (plant breeding and agronomical and chemical approaches) requires research on different levels of organization from molecule to field, and includes all processing systems from farmers to chemists.
  3. Cirak, Cuneyt and ., Ali Kemal Ayan and ., Kudret Kevseroglu, The Effects of Light and Some Presoaking Treatments on GerminationRate of St. John’s Worth (Hypericum Perforatum L. {$<$}i/{$>$} ) Seeds, Pakistan Journal of Biological Sciences, vol. 7, no. 2, pp. 182--186, January 2004. doi: 10.3923/pjbs.2004.182.186.
  4. {Pérez-García}, F. and Huertas, M. and Mora, E. and Peña, B. and Varela, F. and {González-Benito}, M. E., Hypericum Perforatum L. Seed Germination: Interpopulation Variationand Effect of Light, Temperature, Presowing Treatments and Seed Desiccation, Genetic Resources and Crop Evolution, vol. 53, no. 6, pp. 1187--1198, September 2006. doi: 10.1007/s10722-005-2012-3.
    Germplasm conservation of medicinal plants is of increasing interest and, when possible, seed banking is the most efficient system for ex situ conservation of these plant genetic resources. Hypericum perforatum L. (St. John's wort, Guttiferae) is a medicinal plant with evidence of efficacy as an anti-depressant. The aim of this work was to increase knowledge of its seed germination behaviour by studying 68 wild populations. Seed germination tests were carried out at 25/15~°C under a photoperiod of 16-h light/8-h darkness. Final germination percentages were highly variable depending on the accession, ranging from 6 to 98\%. Similarly, germination rate (as expressed by T50 values) varied significantly from 6.1 to 23.0~days. The effect of seed desiccation with silica gel on subsequent germination was also studied. The effect of two other incubation temperatures (15 and 25~°C) and light (photoperiod or darkness) on seed germination was studied in several accessions. Temperature had no significant effect on final germination percentages. However, light significantly increased the germination of most but not all accessions assayed. Seeds from four accessions with low germination percentages were subjected to different presowing treatments that could increase germination: dry heat, hot water and gibberellic acid. Germination was promoted significantly by gibberellic acid in two of the four accessions assayed, but the thermal treatments did not enhance significantly the germination percentages. This study reveals that conclusions based on one population of Hypericum perforatum cannot characterize the germination behaviour for the entire species.
  5. Borthwick, H. A., Retarded Germination in the Seed of Hypericum Perforatum Caused by Calcium, Botanical Gazette, vol. 98, no. 2, pp. 270--282, December 1936. doi: 10.1086/334636.
    1. Germination studies with Hypericum perforatum seed showed that tap water retards germination as contrasted with the germination rate in distilled water. Mixtures of the two containing as little as 10 per cent tap water cause as pronounced germination delay as pure tap water. A relatively short period in tap water, followed by transfer to distilled water, also results in a definite retardation. 2. High alkalinity of the tap water and its ionic constitution are both possible causes of the effect observed. The data show that it is not a question of alkalinity, for similar results can be produced in solutions at or below the neutral point. The effects of the various ions present in tap water indicate that calcium causes the retarded germination. 3. No conclusive explanation is offered as to the method of operation of calcium in delaying germination. The data suggest that it alters the permeability of the coat to water.
  6. Waters, S. J. P. and Pigott, C. D., Mineral Nutrition and Calcifuge Behaviour in Hypericum, Journal of Ecology, vol. 59, no. 1, pp. 179--187, 1971. doi: 10.2307/2258460.
    Three glasshouse experiments were carried out to investigate the calcifuge behaviour of Hypericum humifusum and H. pulchrum, by comparing their growth with that of the lime-tolerant H. perforatum on two acid and two highly calcareous soils. Additions to the soil of phosphate, chelated iron, nitrate, potassium and manganese were made. All treatments were combined factorially. Addition of phosphate alone produced large increases of dry matter yield in all species on the acid soils and in H. perforatum on the calcareous soils, but produced a rather small increase in growth of H. pulchrum and no increase in growth of H. humifusum on the calcareous soils. Addition of chelated iron alone produced large growth increases in H. pulchrum and in H. humifusum on the calcareous soils, accompanied by relief of lime-induced chlorosis and, in H. humifusum, promotion of flowering. Phosphate and chelated iron added in combination showed a strong positive interaction in promoting growth and flowering in H. humifusum on one of the calcareous soils. A preliminary outdoor experiment indicated clearly that additional factors must be considered in natural environments.
  7. Campbell, M. H., Germination, Emergence and Seedling Growth of Hypericum Perforatum L., Weed Research, vol. 25, no. 4, pp. 259--266, 1985. doi: 10.1111/j.1365-3180.1985.tb00643.x.
    Germination of new seeds (1–6 months old) of Hypericum perforation L. was restricted by high temperatures (16h/8h, 20/30°C), darkness and a chemical inhibitor in exudate from the capsule, whereas germination of old seeds (9 years) was only restricted by the inhibitor. The effect of the chemical inhibitor and high temperatures was overcome, respectively, by washing seeds in water and by reducing temperatures to constant 15°C. Calcium in solution from CaCO3 and from three different soils did not prevent the germination of new or old seeds or of seeds collected from five different locations. There were differences in the germination characteristics and dormancy mechanisms of seeds collected from different localities, Restriction of the emergence of seedlings by a covering of {$>$} 2 mm of soil appeared to be due to lack of seedling vigour rather than to lack of light. Seedling growth was much slower than in other pasture species. Thus the requirements for germination of H. perforatum of low temperature and moisture to wash away the chemical inhibitor favour its establishment but the slow growth of its seedling restricts its emergence and renders it extremely susceptible to competition from other plants.
  8. Chen, Chung Li and Tsai, Yuen Jen and Sung, Jih Min, Photoperiod Effects on Flowering and Seed Setting of Hypericum Perforatum, Experimental Agriculture, vol. 46, no. 3, pp. 393--400, July 2010. doi: 10.1017/S0014479710000050.
    The effects of day length extension and night interruption on flowering responses of St John's Wort (Hypericum perforatum) plants were investigated. Field-grown plants were subjected to five different day lengths (11, 13, 15, 17 and 19 h d−1) or night interruption for 70 days. The results indicated that St John's Wort is a long-day plant requiring a critical day length of 15 hd−1 for flowering induction. Both day length extension (duration longer than 15 h) and night interruption, with artificial light (fluorescent lamps that delivered 30 μmol m−2 s−1 photosynthetic photon flux density at plant height) were effective for flowering induction and seed setting. Day length extension or night interruption experienced by the maternal plant also affected the germination responses of the seeds produced. Plants that had received 19 h of day length treatment produced seeds with better germination responses. Night interruption with daylight type fluorescent lamps was also effective for producing relatively high quality seeds, although these seeds had slightly lower germination rates and longer mean germination time than seeds produced under 19 h of day length. Such a night interruption system could be considered for seed production of St John's Wort on a commercial scale.
  9. Butola, Jitendra S and Pant, Shreekar and Samant, S S, Effect of Pre-Sowing Seed Treatments in Hypericum Perforatum L: A High Value Medicinal Plant, Seed Research, vol. 35, no. 2, pp. 205--209, 2007. url: https://www.academia.edu/55185905/Effect_of_Pre_sowing_Seed_Treatments_in_Hypericum_perforatum_L_A_High_Value_Medicinal_Plant.
    Hypericum perforatum L. is a high value vulnerable medicinal plant of the Indian Himalayan Region. Seeds of this species showed poor germination. In present study, of the total 15 pre-sowing treatments tried, 13 significantly (P {$<$} 0.05) stimulated seed-germination, where GA3 (150 mM), KNO3 (150 mM) and NaHClO3 (15 minutes) were found to be most effective. The control showed only 27.8\% germination while it increased highest upto 71.1\% by KNO3 (150 mM). These treatments were also effective in reducing the time required for germination and mean germination time. In view of the low cost and easy applicability of KNO3 and NaHClO3 compared to the expensive and technically complicated plant growth regulators, these chemicals could be used by poor and unskilled farmers of the Indian Himalayan Region for the ex-situ cultivation of H. perforatum.
  10. Semenko, Maksym and Pospelov, Sergii, Technological Aspects of St. John's Wort (Hypericum Perforatum L.) Cultivation, Grail of Science, no. 16, pp. 160--162, July 2022. doi: 10.36074/grail-of-science.17.06.2022.027.
    Despite the widespread raw material base of St. John's wort in nature, there is an urgent need to cultivate it [4,5]. In this regard, there is already some experience in growing crops and developed the basic elements of cultivation technology.
  11. Naghdi Badi, H. and Ziai, S. A. and Mirjalili, M. H. and Ahvazi, M. and Khalighi Sigaroodi, F. and Habibi Khaniani, B. and Farahani, A., Variation in Quantitative Yield and Hypericin Content of St. John’s Wort., Hypericum Perforatum L., Journal of Medicinal Plants, vol. 3, no. 11, pp. 59--67, September 2004. url: http://jmp.ir/article-1-738-en.html.
    St John\’s wort, Hypericum Perforatum L. (Culsiaceae) is an important medicinal plant, which has different bioactive constituents and hypericin (a naphtodianthrone) is one of this compounds. Hypericin has many pharmacological effects such as antidepressant, antiviral and antibacterial, which cause to high production and consumption. It has known that region and ...
  12. Azizi, M. and Omidbaigi, R., Effect of Np Supply on Herb Yield, Hypericin Content and Cadmium Accumulation of St. John's Wort (Hypericum Perforatum L.), Acta Horticulturae, no. 576, pp. 267--271, April 2002. doi: 10.17660/ActaHortic.2002.576.39.
  13. Hevia, F. and Berti, M. and Wilckens, R., Quality and Yield in St. John's Wort (Hypericum Perforatum L.) Harvested in Different Phenological Stages, Acta Agronomica Hungarica, vol. 50, no. 3, pp. 349--358, July 2005. doi: 10.1556/AAgr.50.2002.3.12.
    Two experiments were conducted in the province of Ñuble, Chile during the 1997/98 and 1998/99 seasons with the objective of evaluating the effect of harvesting date on the yield and quality of St. John's wort (Hypericum perforatum L.) in the second year of production. The apical 25 cm of the stem were harvested in the following stages: flower bud, beginning of flowering, full flower and petal drop. A randomized complete block design with four replications was used. The best yield (fresh, dry and threshing weight) and the highest hypericin content were obtained at the petal drop stage. Nevertheless, the results indicate that the best time to harvest St. John's wort is when 10 to 20\% of the flowers are open and the rest are in the bud stage.
  14. Franke, R. and Schenk, R. and Bauermann, U., Variability in Hypericum Perforatum L. Breeding Lines, Acta Horticulturae, no. 502, pp. 167--174, December 1999. doi: 10.17660/ActaHortic.1999.502.26.
  15. Bagdonaitė, Edita and Labokas, Juozas, Morphological Variability of the Field Accessions of Hypericum Perforatum, Vytauto Didžiojo universiteto Botanikos sodo raštai, vol. 11, pp. 8--13, 2006. url: https://vb.gamtc.lt/object/elaba:6223016/.
    Variation of ten phenotypic characters was assessed among fifteen Lithuanian accessions of Hypericum perforatum L. of wild origin as well as Polish cultivar "Topaz" cultivated in the field collection of medicinal plants at the Institute of Botany. High variations were established within accessions in length and width of inflorescence and weight of raw material. It was established that the morphotypes vary very much within the accessions as well. The accessions No. 381, No. 419, No. 423, No. 424, No. 426 and No. 427 are adequate to the cv. "Topaz" and produce fairly long and wide inflorescences with high total yield of medicinal raw material.
  16. Giurgiu, R.M. and Morar, G. and Dumitraș, A. and Vlăsceanu, G. and Dune, A. and Schroeder, F.-G., A Study of the Cultivation of Medicinal Plants in Hydroponic and Aeroponic Technologies in a Protected Environment, Acta Horticulturae, no. 1170, pp. 671--678, July 2017. doi: 10.17660/ActaHortic.2017.1170.84.
  17. Bagdonaitė, Edita and Janulis, Valdimaras and Ivanauskas, Liudas and Labokas, Juozas, Ex Situ Studies on Chemical and Morphological Variability of Hypericum Perforatum L. in Lithuania, Biologija, vol. 53, no. 3, July 2007. url: https://www.lmaleidykla.lt/ojs/index.php/biologija/article/view/754.
    This study describes the variation of hypericin and flavonoid contents in different accessions of Saint John’s Wort, Hypericum perforatum L. Twenty-one Lithuanian wild accessions as well as two cultivars, Polish ‘Topas’ and Russian ‘Zolotodolinskaya’, were studied under the same cultivated field conditions with the latter two used as control. The chemical and morphological investigations were carried out in two-year-old plants. Samples of flowering tops of H. perforatum were collected and analysed for hypericin and flavonoids using high performance liquid chromatography (HPLC) analysis. The results showed that hypericin contents in flowering tops ranged within 0.23–1.24 mg/g; flavonoid contents varied in different accessions as follows: rutin 2.95–17.10 mg/g, hyperoside 0.42–31.13 mg/g, quercitrin 0.16 to 7.52 mg/g, and quercetin 0.37–1.90 mg/g. The results revealed a reliable relation between the contents of hypericin and the morphotypes of H. perforatum. Some of the accessions of H. perforatum are distinguished by higher contents of the secondary metabolites studied if compared with the cv. ‘Zolotodolinskaya’ and ‘Topas’ and could be used for breeding purposes. Keywords: Hypericum perforatum, hypericin, flavonoids, morphotypes
  18. Alan, Ali R. and Murch, Susan J. and Saxena, Praveen K., Evaluation of Ploidy Variations in Hypericum Perforatum L. (St. John’s Wort) Germplasm from Seeds, in Vitro Germplasm Collection, and Regenerants from Floral Cultures, In Vitro Cellular \& Developmental Biology - Plant, vol. 51, no. 4, pp. 452--462, August 2015. doi: 10.1007/s11627-015-9708-7.
    Hypericum perforatum L. (St. John’s wort) is an important medicinal herb and a subject of intensive research for its complex and diverse bioactive chemicals. An in vitro-grown germplasm collection of elite H. perforatum lines, established to provide easy access to physiologically uniform plants, was used for ploidy assessment studies. Germplasm lines were maintained by repeated subculture of shoot tips for over 10~yr with little change in their capacity to produce multiple shoots. Shoots of four of these lines were rooted and grown in the greenhouse to obtain plants to provide anther and filament explants. Culture of explants on a regeneration medium supplemented with 1~mg~L−1 α-naphthaleneacetic acid (NAA) and 1~mg~L−1 6-benzylaminopurine (BA) induced large numbers of calluses and shoots on all explants. Flow cytometric (FCM) analysis of nuclei samples revealed that the nuclear DNA contents of calluses and shoots developed from anther and filament explants of germplasm lines were not significantly different from those of the donor plants. FCM screening of in vitro-maintained germplasm lines in the collection showed that they had similar nuclear DNA amounts and were all tetraploid (2n\,=\,4x). Analysis of seedlings obtained from the original seed source used to derive the germplasm lines showed that \textasciitilde 11\% of them were hexaploid (2n\,=\,6x). Data obtained from FCM screens confirmed the preservation of tetraploidy in in vitro-maintained H. perforatum germplasm and the regenerants obtained from male floral organs. The consistent ploidy of the H. perforatum plants of in vitro origin further supports the usefulness of such technologies to ensure genetic uniformity of medicinal plants over extended periods of culture and may facilitate long-term preservation of their elite clones.
  19. {DANIELA CICCARELLI} and {ANDREA CESARE ANDREUCCI} and {ANNA MARIA PAGNI}, The Black Nodules of Hypericum Perforatum L. Subsp. Perforatum: Morphological,Anatomical, and Histochemical Studies during the Course of Ontogenesis, Israel Journal of Plant Sciences, vol. 49, no. 1, pp. 33--40, January 2001. doi: 10.1560/46Y5-AFWD-TCY0-KGFG.
    Hypericum perforatum L., traditionally used in folk medicine as a therapeutic plant, is today investigated for its antidepressant and antiretroviral activities. This species is characterized by the presence of different types of secretory structures: translucent glands or cavities, black nodules, and secretory canals. The aim of the present work is to characterize the black nodules on both the floral and vegetative parts, morphologically, anatomically, and histochemically. Nodules consist of a cluster of irregularly shaped cells surrounded by a single- or double-layered sheath. Histochemical tests show that the nodules are negative for the presence of lipids, essential oils, sesquiterpene lactones, steroids, and proteins and positive for pectic-like substances, tannins, and alkaloids. Our results show that the inflorescences are richest in nodules and are, there f ore, the best sites for the extraction of the secondary metabolites.