Plantago major
(broad-leaved plantain)

Pictures

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Identity

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Preferred Scientific Name

• Plantago major L.

Preferred Common Name

• broad-leaved plantain

International Common Names

• English: broadleaf plantain; common plantain; greater plantain; plantain; ribgrass; ribwort; white-man's foot
• Spanish: llantén; llantén común; llantén major
• French: grand plantain; plantain majeur
• Portuguese: tanachagem major

Local Common Names

• Germany: Breitwegerich
• Italy: petacciola; piantaggine maggiore
• Japan: onioobako; seiyooobako
• Netherlands: weegbree, groote
• South Africa: broad-leaved ribwort; cart-track plant; indlebe-ka-tekwane; large plantain; larger ribwort plantain; ripplegrass; rippleseed plantain; wild sago
• Sweden: groblad

EPPO code

• PLAMA (Plantago major)

Summary of Invasiveness

Top of page P. major is native to Europe and Asia but is now widely distributed around the world, particularly in temperate, but also tropical parts. It is easily distributed and maintained by anthropogenic activities, particularly soil disturbance and compaction. Its small seeds may be spread as a contaminant. There is a possibility for invasion of naturally disturbed habitats (e.g. riparian) as well as anthropogenically disturbed areas and grasslands.

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Plantaginales
•                         Family: Plantaginaceae
•                             Genus: Plantago
•                                 Species: Plantago major

Notes on Taxonomy and Nomenclature

Top of page A number of subspecies of P. major have been named: for example, dostalii, major, winteri, intermedia and pleiosperma (Penkova, 1986; Akeroyd and Doogue, 1988; Lotz et al., 1990).

Description

Top of page P. major is a glabrous to pubescent perennial with one rosette, leaves ovate to broadly so, abruptly narrowed to petiole usually more or less as long as leaves 1.5-40 cm, entire to weakly toothed. Scapes to 40 cm, not furrowed. Inflorescence a spike up to 20 cm long. Many small flowers subtended by bracts 1-2 mm, ovate, glabrous, brownish white with green keel. Sepals 1.5-2.5 mm, green. Corolla tube ca. 2 mm, glabrous, the lobes ca.1 mm, yellowish white, subobtuse, glabrous. Stamens exserted to 2-3 mm, anthers at first lilac, later dirty yellow. Fruits 2-4 mm. Seeds are two celled. This species is wind pollinated.

Subspecies major: leaves mostly with five to nine veins, usually obtuse at the apex, subcordate to rounded at the base and subentire; capsules mostly with 4-15 seeds; seeds (1)1.2-1.8 (2.10) mm.

Subspecies intermedia: plants usually smaller with much shorter spikes; leaves mostly with three to five veins, usually subacute at apex, broadly cunate at base and +/- undulate toothed near base; capsules mostly with (9)14-25 (36) seeds; seeds (0.6) 0.8-1.2 (1.5) mm.

The morphology of P. major is fairly variable, even within populations and at a small spatial scale of tens of metres for some sites (Lotz et al., 1990). Populations subject to intensive grazing or cutting are generally lower growing and less erect, and variation in growth form has been shown to contain a genetic component (Warwick and Briggs, 1979, 1980).

P. major forms a basal rosette with a compressed stalk and leafless flower stalk. Root contraction has been observed in this species and is related to resistance to treading. The fruit is a capsule opening with an operculum and the seed is mucilaginous and easily transported by cattle or man (Soekarjo, 1992).

Plant Type

Top of page Broadleaved
Perennial
Seed propagated

Distribution

Top of page P. major is a worldwide weed originating from Eurasia (USDA-ARS, 2003). It is distributed widely throughout Europe, North Africa, North and Central Asia and has naturalized throughout most of the world in temperate climates (Clapham et al., 1989); it is also present in some tropical areas.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

Top of page This species is synanthropic (often associated with human activities and landuse such as agriculture, earthworks). It has thus spread from Eurasia to much of the rest of the world, particularly temperate areas. Its pollen is used as an indicator of human settlement from the Neolithicum onwards (Aart and Vulto, 1992a). In the Americas it is known as 'white man's foot' (Mitch, 1987). Godwin (1944) notes the increase in abundance of this species in Denmark's prehistoric pollen record as original forest cover was replaced by cultivated land. The seedbank of this species in Danish arable soil is thought to have decreased significantly over the period 1964-1992 (Jensen and Hjellsson, 1992).

Habitat

Top of page Like other Plantago species, this is a plant of open, well lit (Aart and Vulto, 1992b), often grassy habitats (Stace, 1997) in a range of climates. Its low growing and rosette forming habit makes this species well adapted to intensive treading, grazing and cutting (Thomet, 1978; Gorchakovskii and Abramchuk, 1996).

P. major is found in grass seed mixes for a range of purposes (Kolb and Schwarz, 1983) and therefore it is present in a wide range of seeded grassland habitats. It is tolerant of high nutrient conditions and is found in agriculturally improved grasslands (Aart and Vulto, 1992b). This species is also a plant of man-made habitats such as urban areas and disturbed ground (Bastin and Thomas, 1999).

P. major subsp. intermedia tends to be found in damp, usually saline places near the sea and less often inland (Stace, 1997).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
	Terrestrial – Managed	Cultivated / agricultural land	Present, no further details
		Managed forests, plantations and orchards	Present, no further details
		Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
		Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
		Disturbed areas	Present, no further details	Harmful (pest or invasive)
		Rail / roadsides	Present, no further details	Harmful (pest or invasive)
		Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
	Terrestrial ‑ Natural / Semi-natural	Natural forests	Present, no further details	Harmful (pest or invasive)
		Natural grasslands	Present, no further details	Harmful (pest or invasive)
		Riverbanks	Present, no further details	Harmful (pest or invasive)

Hosts/Species Affected

Top of page P. major is mainly found in grassland.

Biology and Ecology

Top of page Genetics

P. major has a chromosome number of 2n=12 (Stace, 1997). Populations show a high level of genetic structure, with a tendency to form a range of morphological variants specialized to fit particular niches. At least some of this specialization is thought to be due to: 1) the high rate of self-fertilization in the species, which restricts gene flow between variants, and 2) the high chromosome variability of this species. Such specialization and interrupted gene flow contrast with the free gene flow of some other Plantago species which are outbreeding generalists (Sharma et al., 1992; Sharma and Koul, 1995). Alloenzyme variation evidence corroborates this analysis, and shows high variation and low gene flow in P. major (Dijk and Wolff, 1992).

P. major tends to show wide variability across, but uniformity within, populations. Analysis of the inter- and intraspecific variation of chloroplast DNA of four European Plantago species (P. major, P. lanceolata, P. media and P. coronopus) by Hooglander et al. (1993) showed P. major to be most closely related to P. media. A short mutation (70 base pairs) could discriminate between the subspecies major and pleiosperma. Wolff and Morgan-Richards (1998) have also distinguished P. major subspecies using PCR markers.

Physiology and Phenology

P. major seed dormancy is broken by stratification (1-7 days), exogenous gibberellic acid (GA3) and potassium nitrate (Saruhan et al., 2002). Light is normally required for germination (Blom, 1978; Pons and Toorn, 1988; Pons, 1991) and an approximately linear decrease in germination rate with increasing soil depth has been observed (Bliss and Smith, 1985). Imbibition (soaking) and alternating temperature (20°C/30°C) increase germination (Deschenes and Moineau, 1972); ethylene treatment was inhibitory and thereafter germination occurred only in the presence of light (Heydecker et al., 1972). Two- to 5-year-old seed showed a higher germination rate than fresh seed (Blom, 1992). Seeds of this species have been shown to survive passage through the digestive tract of a cow and will germinate if followed by a period of low temperature (5°C) but not when followed by a period of higher temperature (20°C) (Holub and Lhotska, 1991).

Under optimal conditions, P. major shows higher rates of emergence and establishment in uncompacted rather than compacted soils. However the roots of this species have a marked ability to penetrate compacted soils and, under conditions of low moisture, P. major does better in compacted than loose sandy soil (Blom, 1976). Although perennial, P. major may behave as an annual, flowering and setting seeds within 6 weeks of germination. In Britain, flowering normally starts in early June and continues for about 3 months (Sagar and Harper, 1964). A micropropagation protocol for this species has been produced by Mederos et al. (1998) and involves shoot tip culture on modified MS medium.

Reproductive Biology

P. major reproduces mainly by seeds and cannot multiply freely by vegetative means (Holm et al., 1977). The species is wind pollinated, but relies heavily on self-pollination (Sharma et al., 1992; Sharma and Koul, 1995). There may be 3-30 seeds per capsule and a seed production of 14,000 seeds per plant per year has been recorded (Holm et al., 1977). Sharma and Koul (1995) and Sharma et al. (1992) describe the reproductive strategy of this species as making a greater investment in female rather than male reproductive components and relying heavily on self-fertilization. Sagar and Harper (1964) note that seed is set rapidly after fertilization but is frequently not dispersed from the capsules until the following year.


Environmental Requirements

There is some evidence that P. major is more shade tolerant than other members of the genus (Toorn and Pons, 1988). There is also evidence that this species has a higher soil moisture requirement than some other Plantago species (Blom, 1976). Stoutjesdijk (1992) indicates that P. major tolerates a narrower range of temperature, solar radiation and humidity than other Plantago species such as P. lanceolata. Sagar and Harper (1964) note that P. major occurs on a wide range of soil types in Britain, being absent only from extremely acid peats and mountain grasslands.

P. major is particularly resistant to trampling and compaction (Engelaar et al., 1993; Engelaar, 1995; Gorchakovskii and Abramchuk, 1996). Zelikov and Psonnova (1961) and Kolb and Schwarz (1983) found it inhabited soil with a mean density of 1.42 g/cm3, and its abundance has been positively correlated with soil compaction (Crawford and Liddle, 1977; Aspinall and Pye, 1987). Thomet (1978) and Holeksa and Holeksa (1987) found P. major to be an indicator of excessive treading in permanent pastures. A number of studies show this species to be capable of withstanding considerable foot traffic and it is often an important component of well worn turf (Montacchini and Siniscalco, 1982).

Associations

Sagar and Harper (1964) provide detailed lists of plants associated with P. major in the British Isles.

Rainfall

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Soil Tolerances

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Soil drainage

• free
• impeded
• seasonally waterlogged
• seasonally waterlogged

Soil texture

• heavy
• light
• medium

Notes on Natural Enemies

Top of page Sagar and Harper (1964) provide a long list of natural enemies including invertebrates, fungi and viruses but imply that these have much less influence than livestock management.

Means of Movement and Dispersal

Top of page Natural Dispersal (Non-Biotic)

The small seeds are dispersed by the wind.

Vector Transmission (Biotic)

The seeds are mucilaginous and are dispersed by humans and animals (Soekarjo, 1992).

Accidental Introduction

Because of the small size of its seeds, P. major may be introduced as a contaminant of agricultural produce.

Plant Trade

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Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Leaves
Stems (above ground)/Shoots/Trunks/Branches
Wood

Impact Summary

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Impact

Top of page P. major has been described as an agricultural, pastoral and environmental weed competing with other plants for light, water and nutrients and replacing preferred vegetation. P. lanceolata and P. major have together been reported as weeds in over 50 countries affecting a wide range of crops (Holm et al., 1977). It is a field rather than field margin weed, although it colonizes disturbed margins (Kress, 1988).

In the UK it affects the majority of local authority owned sports turf (Raikes et al., 1994). In Prince Edward Island, Canada, it is present in 80% of cereal fields with a mean density of over 14 plants per m² (Thomas and Ivany, 1990).

Risk and Impact Factors

Top of page Invasiveness

• Invasive in its native range
• Proved invasive outside its native range
• Highly adaptable to different environments
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Highly mobile locally
• Has high reproductive potential

Impact outcomes

• Negatively impacts agriculture
• Negatively impacts tourism
• Reduced amenity values

Impact mechanisms

• Competition - monopolizing resources

Likelihood of entry/control

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect as a commodity contaminant
• Difficult/costly to control

Uses

Top of page P. major has medicinal properties and is a popular Chinese medicine. It contains phenylethanoid glycosides. It is used to treat inflammation (Nunez Guillen et al., 1997), gastritis and peptic ulcers (Aye et al., 1996b), leishmanial ulcers (Franca et al., 1996), to reduce pain (Nunez Guillen et al., 1997), and as an antidiarrhoeal agent (Heinrich, 1998). It is also used in the treatment of dermatological conditions (Brown and Dattner, 1998), common cold, viral hepatitis (Chiang et al., 2002), and has expectorant, cicatrizant and astringent properties (Ramos et al., 2002). Aqueous extracts of P. major have a significant effect on aspirin-induced ulceration in rats (Aye et al., 1996b), some effects on pain and inflammation (Nunez Guillen, 1997; Aye et al., 1996a). Navarro et al. (1998) report P. major to be effective at controlling dental plaque and gingivitis. Extracts of this species show some degree of antibacterial activity (Holetz et al., 2002).

P. major is also used as fodder (Fogelfors, 1984). It is higher in trace elements than pasture grasses and therefore can be considered a useful pasture component (Trzaskos, 1996). It is palatable to sheep (Barcsak and Kispal, 1984).

P. major is cultivated as an ornamental in South Africa (Wells et al., 1986).

Uses List

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Medicinal, pharmaceutical

• Traditional/folklore

Similarities to Other Species/Conditions

Top of page P. major is distinguished from other Plantago species by its long-petioled leaf blade and many-seeded capsule that opens around the fruit above the base, and the angular seeds which are marked with thread-like ridges and bear a light-coloured hilum (Holm et al., 1977).

Prevention and Control

Top of page Cultural Control

Flooding and trampling regimes aimed at weed control have not been successful against P. major (Engelaar and Blom, 1995). Similarly, a four-course crop rotation (oats, clover, winter wheat, faba beans), using minimal weed control for 4 years, resulted in large increases in weed biomass and surface and subsurface seed bank of P. major (Hill et al., 1989).

Mechanical Control

Mechanized spraying of hot water (85-95°C) in an orchard required two to three repeat sprays for good control of this and other species (Kurfess and Kleisinger, 2000).

Whilst microwave radiation has been tested as a weed control measure, it required more than 32 h of treatment (at 2.45 Ghz, 6 kW) for complete control of P. major in pot experiments. Higher frequencies (13.5 MHz, 50 kW) were effective against some erect weeds but not P. major (Kunisch et al., 1992).

Chemical Control

Treatment with chlorthal-dimethyl + naptalam resulted in successful control in Cucumis crops (Himme et al., 1984). In winter and spring cereals, glyphosate efficiently controlled weeds, including P. major, by pre-harvest application and on stubble after harvesting (Ciuberkis and Petraitis, 1998).

Amitrole or paraquat with diuron failed to control this species in orchards (Himme and Stryckers, 1975a, b), and it is resistant to lenacil (Stryckers and Himme, 1974) and methazole (Tasmanian Department of Agriculture, 1975). In a long-term repeated spraying regime using diuron, P. major was the first weed species to gain resistance (Bulcke et al., 1989).

P. major is controlled less easily with chemicals than are other turf weeds (Schery, 1974). In Festuca rubra / Poa pratensis turf, MCPA or 2,4-D applied in July-August (in northwest USA) provided excellent control of this species (Ebdon and Jagschitz, 1981). In Festuca arundinacea / Poa pratensis turf, the best control of P. major was obtained with 2,4-D (Wehner et al., 1981). In turf close to ornamental shrubs, dicamba + 2,4-D was effective (Hendricks et al., 1983). Neal and Mascianica (1988) found that quinclorac and triclopyr provided adequate control of this species in turf. However, Neal (1990) found 2,4-D and clopyralid + triclopyr were effective treatments but not quinclorac, chlorflurenol or dicamba. MCPA ± mecoprop + dicamba or clopyralid ± triclopyr or chlorflurenol ± clopyralid gave excellent control of Plantago spp. in turf, including P. major (Sawyer and Jagschitz, 1988).

Heavy applications of glyphosate to pasture increased the abundance of P. major (Grabowski, 1990). Vidme (1973) recommends that weed control by herbicide (such as 2,4 D salts or esters) in grassland is combined with heavy nitrogen applications for maximum effect.

Biological Control

P. major has been identified as a priority weed for bioherbicide research of lawn weeds (Gadoury and Watson, 1987).

Integrated Control

This weed was best controlled in rainfed rice (Nagaland, India) by a combination of chemical (2,4-D, butachlor and fluchloralin at 1 day after transplanting (DAT); thiobencarb at 1 and 7 DAT) and hand-weeding (at 25 and 45 DAT; Sharma et al., 1994).

References

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Aart van der PJM; Vulto JC, 1992. General ecology. In: Kuiper PJC, Bos M, eds. Plantago: a Multidisciplinary Study. Ecological Studies, Vol. 89. Berlin, Germany: Springer Verlag, 6.

Akeroyd JR; Doogue D, 1988. Plantago major L. subsp. intermedia (DC.) Arcangeli (Plantaginaceae) in Ireland. Irish Naturalists' Journal, 22(10):441-443

Andreasen C; Stryhn H; Streibig JC, 1996. Decline of the flora in Danish arable fields. Journal of Applied Ecology, 33(3):619-626; 44 ref.

Aspinall RJ; Pye AM, 1997. The effect of trampling on limestone grassland in the Malham area of North Yorkshire. Journal of Biogeography, 14(2):105-115.

Aye T; Mu MSM; Tin M; Win M, 1996. Anti-oedema activity of Nyctanthes arbor-tristis L., Curcuma longa L. and Plantago major L. Myanmar Health Sciences Research Journal, 8(1):36-40.

Aye T; Mu MSM; Win M; Tin M; Su SH, 1996. The anti-ulcerogenic activity of Plantago major Linn. Myanmar Health Sciences Research Journal, 8(2):74-77.

Barcsak Z; Kispal T, 1990. Palatability examination of grasses. In: Gaborcik N, Krajcovic V, Zimkova M, eds. Soil Grassland Animal Relationships. Proceedings of 13th General Meeting of the European Grassland Federation, Banska Bystrica, Czechoslovakia, June 25-29, 1990, Volume 2. Banska Bystrica, Czechoslovakia: Grassland Research Institute, 281-284.

Basaran AA; Ceritoglu I; Undeger U; Basara N, 1997. Immunomodulatory activities of some Turkish medicinal plants. Phytotherapy Research, 11(8):609-611.

Bastin L; Thomas CD, 1999. The distribution of plant species in urban vegetation fragments. Landscape Ecology, 14(5):493-507; 43 ref.

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Bekmansurov MV, 1996. Investigation of greater plantain (Plantago major L.) consortia. Russian Journal of Ecology, 27(2):148-149.

Bliss D; Smith H, 1985. Penetration of light into soil and its role in the control of seed germination. Plant, Cell and Environment, 8(7):475-483.

Blom CWPM, 1976. Effects of trampling and soil compaction on the occurrence of some Plantago species in coastal sand dunes. I. Soil compaction, soil moisture and seedling emergence. Oecologia Plantarum, 11(3):225-241

Blom CWPM, 1978. Germination, seedling emergence and establishment of some Plantago species under laboratory and field conditions. Acta Botanica Neerlandica, 27(5/6):257-271

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Bottcher W, 1993. Effects of intensive farming on the landscape of a river meadowland in the Saxon hilly country. Archiv fur Naturschutz und Landschaftsforschung, 32(3):209-225

Brown DJ; Dattner AM, 1998. Phytotherapeutic approaches to common dermatologic conditions. Archives of Dermatology, 134(11):1401-1404.

Bulcke R; Himme M van; Stryckers J, 1989. Shifts in the flora by long-term repeated herbicide applications in a bush apple plantation. Proceedings of the 4th EWRS symposium on weed problems in Mediterranean climates. Vol. 1. Problems of weed control in fruit, horticultural crops and rice, 121-128

Chiang LC; Chiang W; Chang MY; Ng LT; Lin CC, 2002. Antiviral activity of Plantago major extracts and related compounds in vitro. Antiviral Research, 55(1):53-62.

Chua KS; Tan HTW; Turner IM, 1994. The angiosperm flora of Singapore. Part 3. Plantaginaceae. Gardens' Bulletin (Singapore), 46(2):103-107; [1 fig.]; 9 ref.

Ciuberkis S; Petraitis V, 1998. Application of Roundup to hasten the ripening of winter and spring cereals and to control the weeds. Z^hacek~emdirbyste^dot over~, Mokslo Darbai, 62:63-75; 12 ref.

Clapham AR; Tutin TG; Moore DM, 1989. Flora of the British Isles. Cambridge, UK: Cambridge University Press.

Coste I, 1970. Contribution to the study of a Molinio-Arrhenatheretea Tx. (1937) 1970 in the Locva Mountains (southwestern Romania). Revue Roumaine de Biologie, Biologie Vegetale, 24(1):17-26.

Crawford AK; Liddle MJ, 1977. The effect of trampling on neutral grassland. Biological Conservation, 12(2):135-142.

Cristea V; Groza G, 1983. Contributions to studies of the vegetation of the hills of 'Batrinu' - Vadu Crisului municipalitu (Bihor district). Contributii Botanice Universitatea 'Babes-Bolyai' din Cluj Napoca, 137-143.

Deschenes JM; Moineau D, 1972. The conditions required for germination in four Quebec weeds. Naturaliste Canadien, 99:103-114

Dijk H van; Wolff K, 1992. Allozyme variation and genetic structure in Plantago species. Plantago: a multidisciplinary study [edited by Kuiper, P. J. C.; Bos, M.] Berlin, Germany; Springer-Verlag, 184-192

Dunst RM; Pool RM, 1985. Preemergence annual weed control in a New York vineyard. Proceedings, 39th annual meeting of the Northeastern Weed Science Society., 211-213.

Ebdon JS; Jagschitz JA, 1981. Evaluation of herbicides for broadleaf weed control in lawn turf. Proceedings, Northeastern Weed Science Society, 1981., Volume 35:274-279

Engelaar WMHG; Blom CWPM, 1995. Effects of flooding and trampling on the performance of river foreland species of Rumex and Plantago. Acta Botanica Neerlandica, 44(3):225-245

Fogelfors H, 1984. Useful weeds? Part 14. Lantmannen, 105(16):43

Forcella F, 1992. Invasive weeds in the northern Rocky Mountains. Western Wildlands, 18(2):2-5

Franca F; Lago EL; Marsden PD, 1996. Plants used in the treatment of leishmanial ulcers due to Leishmania (Vannia) braziliensis in an endemic area of Bahia, Brazil. Revista da Sociedade Brasileira de Medicina Tropical, 29(3):229-232.

Gadoury H; Watson AK, 1987. Biological control of lawn weeds. Cahier des Journees Horticoles Ornementales, III:9-15.

Glen HF, 1998. Investigation of the antiinflammatory activity of liquid extracts of Plantago lanceolata L. FSA contributions 12: Plantaginaceae. Bothalia, 28(2):151-157; 26 ref.

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Gorchakovskii PL; Abramchuk AV, 1996. Grazing tolerance of the vegetation of dry meadows. Russian Journal of Ecology, 27(5):321-325.

Grabowski K; Nowicki J; Grzegorczyk S; Benedycki S, 1990. Improvement of productivity of degraded grasslands by means of direct-into-sod sowing. Soil-grassland-animal relationships. Proceedings of 13th general meeting of the European Grassland Federation, Banska Bystrica, Czechoslovakia, June 25-29, 1990 [edited by Gaborcik, N.; Krajcovic, V.; Zimkova, M.] Banska Bystrica, Czechoslovakia; Grassland Research Institute, 355-358

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Haland A, 1989. Nitrogen, potassium and lime for permanent pasture. Effects on botanical composition and yield. Norsk Landbruksforsking, 3(4):225-232

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Hill NM; Patriquin DG; Kloet SP van der, 1989. Weed seed bank and vegetation at the beginning and end of the first cycle of a 4-course crop rotation with minimal weed control. Journal of Applied Ecology, 26(1):233-246

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Holeksa J; Holeksa K, 1987. Plant communities of trampled sites in the Babia Gora National Park (Western Carpathians). Fragmenta Floristica et Geobotanica, 31-32(1-2):247-259.

Holetz FB; Pessini GL; Sanches NR; Cortez DAG; Nakamura CV; Dias Filho BP, 2002. Screening of some plants used in the Brazilian folk medicine for the treatment of infectious diseases. Memo^acute~rias do Instituto Oswaldo Cruz, 97(7):1027-1031; 24 ref.

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Holub M; Lhotska M, 1991. Influence of bovine digestive tract on germination of diaspores of selected plant species - II. Biologia Bratislava, 46(1):81-87.

Hooglander N; Lumaret R; Bos M, 1993. Inter-intraspecific variation of chloroplast DNA of European Plantago spp. Heredity, 70(3):322-334

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