Plantago major L., Greater Plantain

Account Summary

Growth form and preferred habitats

A large and long-persistent soil seed bank enables this all-too-familiar, yet phenotypically extremely variable, weedy, rosette-forming perennial (or annual or biennial), to germinate at any time during the growing season (with a peak flush in late May) (Hawthorn 1974). Irrespective of seasonality, it can very quickly colonise open, short-turf grassland, or bare, disturbed, compacted, wet or dry, very fertile, heavy soil types. Germinating seedlings show effective root penetration in compacted soil (Blom 1979) and the species tends to be especially frequent in heavily trampled ground, including around gateways and along lanes and roadside verges. Experimental work has found that the primary roots from small seeded Plantago species, such as P. coronopus (Buck's-horn Plantain) and P. major subsp. major, are more capable of penetrating the surface of the substrate than those of relatively large seeds such as P. lanceolata (Ribwort Plantain) and P. media (Hoary Plantain) (Blom 1978).

P. major is capable of forming almost pure stands occupying the zone closest to maximum disturbance, although it often associates with Matricaria discoidea (Pineappleweed) and Poa annua (Annual Meadow-grass). Even in closed turf, if P. major manages to colonise any gaps, it will quickly establish and become very persistent as gardeners know to their cost. It can become a real nuisance in lawns (Sagar & Harper 1964).

The primary roots of the seedling are replaced early by a fibrous, adventitious root system from a very short (1 cm), woody taproot or caudex that eventually represents the overwintering food storage organ of the mature, established plant. In P. major, some of the later roots are contractile and help to keep the growing point at the soil surface where it is protected (Holm et al. 1977). In the spring, established plants quickly develop a rosette of spirally arranged, broad, ovate basal leaves. The broad leaves are 5-10 cm long, have an entire or obscurely toothed margin and have 5-7 parallel ribs converging into a rather long petiole. Rapid growth and long leaf petioles allow the plant to produce its characteristic wide-spreading leaf rosettes that establish and occupy valuable surface space. The individual plant can then tenaciously hold its ground against almost all other species, forming small or larger aggregations of offspring around it (Holm et al. 1977).

P. major is generally deep-rooted and can tolerate a range of soil reaction from moderately acid to calcareous (Grime et al. 1988, 2007). The mature root system of P. major may exceed 80 cm in depth and 80 cm in diameter (Kutschera 1960), which enables the species to survive drought conditions. Being resistant to both water-logging and drought and tolerant of half-shade, Greater Plantain appears in a wide variety of lakeshore habitats and even to a limited extent in open areas in woodland in Fermanagh. Generally, it avoids only extremely acid peat or mineral soils, mountain grassland, deep shade, or sites that are continually wet during the growing season (Hawthorn 1974).

The established strategy of Greater Plantain is categorised as intermediate between Ruderal and a balanced C-S-R species (ie Competitor-Stress-tolerator and Ruderal). It is essentially a lowland species of heavily disturbed, fertile, productive vegetation, but it can occur and be widespread up to around 450 m (Grime et al. 1988, 2007).

When growing in exposed places, P. major overwinters as a bud on the woody rootstock at or near ground level. On the other hand, when growing in herbage, an above ground leaf rosette is maintained in all but the most severe winters (Bates 1935).

As Sagar & Harper (1960) point out, many aspects of the life-cycle of P. major suit it for life as an annual species, for example, its self-compatibility, rapid establishment from seed, quick flowering and equally quick maturation of seed. Nevertheless, P. major is capable of persisting as a true and potentially long-lived perennial.

Flowering reproduction

Flowering begins in early June and continues for about four months with a peak in July. Each rosette produces one to many short, leafless scapes bearing the characteristic, cylindrical spike inflorescence that can vary between 1-30 cm long (Holm et al. 1977). The flowers are greatly reduced, wind-pollinated and capable of full self-fertilization. The individual flowers are insignificant in appearance, scentless and without nectar. Seed set is rapid after fertilization, some fruits maturing 2-3 weeks after flowering (anthesis). The species normally produces abundant seed each year and the seed, which is always viable, is small, hard and flattened (Sagar & Harper 1964; Hawthorn 1974).

While some early formed capsules shed seed in autumn, frequently capsules do not disperse their seed until the following spring or summer. Although the capsule splits in the circumscissile manner characteristic of the genus, ie opening around its equator, there is no active dehiscence of seed from the capsule after it splits open. Thus seeds must be knocked off the plant by animals, wind or water. If the seeds are not taken off, they remain attached to the spike until it rots at the base and falls to the ground the next year (Holm et al. 1977).

The number of seeds per capsule varies from 3-28 and the estimated seed production for a large plant is reckoned at c 14,000 per year (Kerner 1896). The seed shape varies with the number contained in individual capsules, a clear indication of the degree of variability within the species, in that a character usually so very conservative can be modified in P. major.

While the main period for seed germination is in spring (peak in May), seeds also germinate intermittently throughout the growing season (Hawthorn 1974) from a large, persistent soil seed bank (Thompson et al. 1997). Seeds of P. major gave 10% germination after burial for 40 years (Crocker 1938).

Despite the purple colour of the anthers, the spikes in full bloom are much less conspicuous than they are in fruit. Since the seeds are attractive to birds, which aid their dispersal, the species bears several English common names that emphasise this property, such as 'Birdseed' and 'Bird's Meat' (Salisbury 1964). In water, an adhesive layer of mucilage on the seed coat (testa) of P. major swells to increase the seed size by about 40%. The ability to imbibe water increases the stickiness of the seeds to soil particles, feathers and fur and greatly enhances the potential for dispersal (Hawthorn 1974).

Variation

Plants vary greatly in size and appearance according to the environmental conditions, ie it is phenotypically very plastic (Salisbury 1964). Infraspecific variation is extensive in P. major and has resulted in a number of lengthy subdivisions into subspecies, varieties and forms (Hawthorn 1974). Sagar & Harper (1964) described Greater Plantain as, "certainly one of the most variable species in the British flora". Sell & Murrell (2007) subdivide P. major into two subspecies, subsp. major and subsp. intermedia, and list four varieties of the latter.

Although subspecific variation has not been sought out in the VC survey, the Fermanagh plant is undoubtedly subsp. major which has cordate leaves with 5-9 veins. In comparison, the leaves of subsp. intermedia have 3-5 veins and the leaf blade narrows to the petiole (Garrard & Streeter 1983).

Hybrids

No hybrids have ever been reported and attempts to produce them artificially have failed (Sagar & Harper 1964).

Vegetative reproduction

Unlike P. lanceolata (Ribwort Plantain) and P. media (Hoary Plantain), P. major does not reproduce freely by vegetative means. While reproduction is primarily by seed, about 20% of plants in trampled laneways and in recently disturbed soil also produce ramets or offsets from buds on the persistent woody crown, undoubtedly in response to damaged tissues. No plants in pasture produce ramets. The ramet remains attached to the parent plant for two to three years, after which the bond breaks and the ramet initiates root development. The daughter plants produced in this way are immediately capable of flowering and seed production (Hawthorn 1974).

Economic importance as a weed

As with P. lanceolata, the success of P. major has been very closely linked to man's activities and especially with agriculture, for a very long time (Holm et al. 1977). In local situations, P. major can reduce the early growth of arable cereal crops such as wheat and oats. While originally P. major and P. lanceolata were temperate zone plants, they have both entered the warmest regions of the world to become pests of almost all tropical crops. They are unique in that more than 50 countries report them as weeds in 26 crops that are as different as alfalfa, coffee, temperate orchards, rice, pastures, cereals, onions, cotton and small fruits. They are most serious competitors in cereals and are common in arable fields of sugar producing areas. Their seeds frequently contaminate small-seeded legume seed-stocks (Holm et al. 1977).

In pasture (and in lawns), P. major plants benefit from occasional mowing as the tall vegetation of competing species is temporarily removed. As expected, P. major grows best in hay pasture where grazing and trampling are prevalent (Hawthorn 1974). The most important factor governing the distribution of P. major around the world is the disturbance of an area by man or animals (Holm et al. 1977).

P. major also hosts a number of viral, fungal and insect pathogens (Hawthorn 1974).

Fossil record

Continuous pollen and macrofossil records from the Weichselian and Flandrian periods prove that P. major is a species of ancient native status in Britain at least (Godwin 1975).

Fermanagh occurrence

Very common throughout Fermanagh on disturbed ground including heavily grazed and/or thoroughly trampled grassland, margins of cultivated ground, lawns, gateways, farmyards, car parks, tracksides, roadsides and waste ground. P. major has been recorded in 447 Fermanagh tetrads, 84.7% of the VC total, which means that it ranks 28th most widespread vascular plant in terms of tetrad frequency.

British and Irish occurrence

In suitably disturbed sites, P. major is extremely widespread and virtually ubiquitous throughout both B & I; in the New Atlas map there are very few hectads where it has not been recorded, most of them probably representing high ground above 630 m (G.M. Kay, in: Preston et al. 2002).

European and world occurrence

The species belongs to the Eurasian Wide-temperate phytogeographical element and is very widespread across Europe, N Africa and N & C Asia. It has been very widely spread by man across both hemispheres, eg to N America where it occurs parallel to indigenous related taxa (Hultén & Fries 1986, Map 1725).

Uses

The fact that plantains frequently grow on paths and around gateways where they get trampled upon, crushed and damaged suggested to herbalists that the plant was so powerful, tough and elastic it must heal crushing, tearing and bruising wounds in humans (Grigson 1955, 1987). Dioscorides wrote that Plantain was a vulnerary (wound-healing herb) and was good in treating ulcers, sores and other forms of wounds or blemishes, including burns. In time, Plantain (probably both P. major and P. lanceolata) became herbs of medico-magical power and divination, the root worn around the neck, for example, as an amulet against the King's Evil.

Grieve (1931) describes P. major as being refrigerant, diuretic, deobstruent and somewhat astringent. It has been used in treating skin inflammation, malignant ulcers, intermittent fever and as a vulnerary. The fresh leaves were also used to staunch bleeding, applied externally as a poultice (Grieve 1931). Additionally, the leaves were used to treat piles, varicose veins, festering wounds and to draw out pus from infected wounds or swellings. It was also considered useful for relieving pain, as in stings. In Ireland, plantain juice was used for coughs and the plant was put to very many other medicinal uses including gout and for treating sore eyes (Allen & Hatfield 2004).

Names

No less than 31 English common names are listed by Grigson (1955, 1987), at least six of which refer to birds, since they are reputed to help disperse the seeds, eg 'Canary Flower', 'Canary Food', 'Lark Seed'. Other names refer to the supposed healing powers of the herb, eg 'Healing Blade', the Scottish 'Slanlis' and the Irish equivalent 'Slanlus', meaning 'healing herb' (Grigson 1955, 1987).

Threats

None.

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