The effects of cutting and asulam on numbers of frond buds and biomass of fronds and rhizomes of bracken Pteridium aquilinum

The effects of cutting and the herbicide asulam for the control of a dense uniform stand of bracken Preridium aquilinum were investigated experimentally over a 6 —yr period in Norfolk, England. Bracken is a particularly difficult species to control by either mechanical or herbicidal means because of its extensive and complex rhizome network, containing large reserves of dry matter and a large number of frond buds. Cutting bracken in the summer removed dry matter from the system and reduced frond vigour but did not reduce the number of frond buds to a critical level. Whereas the first few cuts substantially reduced the standing crop of fronds, relatively stable standing crops of fronds were observed in future years despite continued cutting. Asulam caused a 99% reduction in the standing crop of fronds in the year after spraying, but in the absence of further treatment, fronds recovered to pre-spray levels after 6 yr. Asulam caused severe localised damage to frond buds and rhizome apices, but otherwise the rhizome network remained intact. New sections of frond bearing rhizome (short shoots) developed on storage rhizome (long shoots) deep in the soil of both the asulam plots and those cut.     

The effects of control on the biomass, carbohydrate content and bud reserves of bracken (Pteridium aquilinum (L.) Kuhn), and an evaluation of a bracken growth model

This paper examines the initial effects of bracken control on frond numbers and biomass, and the biomass, carbohydrate reserves and bud densities of bracken stands cut once per year, twice per year, subject to a single application of asulam or left untreated. The seasonal dynamics of these parameters are displayed; carbohydrate and biomass are both removed from the rhizome system to produce frond tissue, and are replenished at the end of the growing season. Asulam application reduced densities of both active and dormant buds, and both frond biomass and density. It did not significantly reduce rhizome biomass or carbohydrate reserves in the two years after treatment. Cutting, either once or twice per year reduced both rhizome biomass and rhizome carbohydrate reserves, as well as bud densities, though the latter were reduced in proportion to biomass. Cutting twice a year reduced the production of fronds, both in numbers and biomass. The collected data were used to evaluate a model of bracken growth, and subsequently to improve estimates of some of the model parameters. The model simulations of control treatments were compared to field data. The effects of cutting once per year and spraying with asulam were predicted accurately, but the bracken stand was more resilient to cutting twice per year than would be expected from model predictions. The combination of cutting and spraying is discussed as a potential tool in land management and the deficiencies of the model are discussed in relation to the need for future research into the biology of bracken.     

The effects of cutting pre-treatments on the performance of asulam on bracken (Pteridium aquilinum)

The performance of asulam was assessed from a field experiment in which bracken (Pteridium aquilinum) was subjected to a variety of cutting pre-treatments prior to spraying with the herbicide. Both cutting bracken twice in the year prior to asulam application, and spraying frond regrowth 10 wk after a single mid-June cut, improved the performance of asulam. Summer cutting removes nutrients and dry matter from the system and reduces frond height, but increases frond density and the number of active buds on the rhizome system. The enhanced efficiency of asulam on pre-cut plots may be due to any one of these factors, or a complex interaction between some (or all) of them. In areas of tall bracken a cutting pre-treatment has the important practical advantage of reducing the height of the canopy, making the subsequent use of ground spraying equipment easier.     

Seasonal variations in the carbohydrate content of bracken

The amounts of dry matter, mobile carbohydrate and reserve carbohydrate have been determined in bracken fronds and buds, frond-bearing rhizomes and storage rhizomes over a period of 13 months. Dry matter and reserve carbohydrate levels are closely linked, especially in the rhizomes, falling from May to the beginning of July and rising until complete frond death at the end of September; but variations occur between frond-bearing and storage rhizomes. The carbohydrate reserve of frond-bearing rhizome is rapidly exhausted by the developing fronds and further carbohydrate is drawn from the storage rhizomes, but a concurrent fall in mobile carbohydrate suggests that the rate of conversion of reserve carbohydrate is slower than the rate of translocation to the fronds. By August, there is an accumulation of carbohydrate in the frond-bearing rhizome, which is followed by a fall in September and this again suggests low enzyme activity in the storage rhizome, but also that sink strengths in bracken are in the order (1) developing buds, (2) storage rhizome, (3) frond-bearing rhizome.
The responses of bracken to cutting and herbicide-application are discussed in relation to sink strengths and a low level of enzyme activity in the storage rhizome.     

The effect of glyphosate on frond regeneration, bud development and survival, and storage rhizome starch content in bracken

In addition to very effectively suppressing bracken frond regeneration for two years at levels of 2.24 kg a. i./ha and above, glyphosate had considerable effects on the rhizome system. Dry weight of frond-bearing rhizomes, number of living apices and developed buds, number and viability of dormant buds and starch content of storage rhizomes were all markedly reduced. The effects on dormant buds and starch content are particularly important when considering frond regeneration and as a consequence of these, glyphosate is likely to give long lasting control of bracken. Low levels of glyphosate (0.02 and 0.07 kg a. i./ha) did not stimulate bracken growth, while the addition of NH₄SCN apparently inhibited the action of the sub-lethal rate of 0.56 kg a. i./ha.     

Efficacy of bracken (Pteridium aquilinum (L.) Kuhn) control treatments across a range of climatic zones in Great Britain

Bracken control field experiments were conducted at six locations across Great Britain. The effects of various cutting and herbicide management regimes upon the seasonal dynamics of bracken fronds and rhizomes were examined over a three year period. This enabled a national overview towards bracken control to be constructed. Initially, spraying with asulam was the most effective treatment in reducing frond biomass and density but was least effective in reducing rhizome biomass. Differential reductions in rhizome biomass were observed in relation to cutting frequency, with cutting twice yearly giving superior control. The national trends confirmed a number of previous observations from independent single-site studies; however, others were contradicted. Comparable management options were ranked consistently between the current and former investigation, however, the initial importance of differences in cutting frequencies did not agree between studies. This multiple-site study improves understanding of the consequences of a national bracken control programme by reducing the influence of confounding site-specific factors, and recommendations for the most appropriate bracken control techniques are made. The extent to which individual sites reflect a national trend in response to bracken control is considered and sites are compared. The hierarchy of treatments identified at the national scale was found to apply generally within individual sites. However, several responses which proved significant at the countrywide level were not so clearly defined at the site scale. Cutting once yearly was the only management regime which appeared to give different bracken control between sites. All other treatments gave similar responses between sites. This result was found in the Scottish Borders, during the second year of control, when frond biomass and density (relative to untreated plots) were greater than that recorded at other sites. This contrast was not found in subsequent monitoring. In terms of rhizome biomass depletion, poorer control was achieved following cutting once yearly at the northern sites (Mull, Scottish Borders, Lake District) compared with the southern sites (Clwyd, Breckland, Devon). The implications of experimental results are discussed in relation to increased cost effectiveness of national bracken control programmes.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

Evaluation of a bracken (Pteridium aquilinum (L.) Kuhn) growth model in predicting the effects of control strategies across a range of climatic zones in Great Britain

A nationwide bracken control experiment provided data over a three-year period for testing the accuracy of a bracken growth model (BRACON). Objective assessments of model validity identified the model as a reasonably accurate predictor of bracken stand dynamics given the range of environmental conditions currently prevailing in Great Britain, in relation to (1) cutting regimes and (2) spraying asulam. Ranking of treatment efficacy at individual sites was closely reflected in model predictions. Predicted response of the rhizome system to cutting treatments underestimated bracken resilience. Failure to consider the consequences of frond regrowth in the latter portion of the growing season as a means to offsetting energy loss was identified as a potential explanation for this discrepancy. The role of the model as a practical management tool is discussed with particular reference to relating model predictions, based upon generalised environmental data (40 km plusmn; 40 km grid cells), to bracken management at individual sites.     

Control of bracken

Mechanical methods of bracken control are now being replaced by the use of chemicals. Trials indicate the value of asulam and glyphosate in reducing frond numbers of bracken when sprayed in late July or early August in the west of Scotland. There is still a need to find a bracken eradicant chemical rather than a control chemical. Research is also required to investigate the results of removing the bracken cover on the ecology of the treated areas. The effects of sudden exposure of the hitherto protected grass to extremes of climate are not known.     

Recovery of Moorland Vegetation after Aerial Spraying of Bracken (Pteridium aquilinum (L.) Kuhn) with Asulam

Bracken (Pteridium aquilinum) is a major weed of seminatural vegetation in Great Britain, as in many other countries. As a consequence, large areas have been subject to control with the intention of restoring the former vegetation. The use of aerial spraying with asulam, a narrow spectrum, carbamate herbicide, has become a common method of control. However, its long‐term efficacy has not been assessed in terms of either controlling the bracken or in producing suitable restored habitats. This study undertook such an assessment by comparing the results of a 2002 survey of the sites subject to bracken control by aerial spraying of asulam with previous (1990/1991 and 1994) surveys of the same sites. This showed that a single application of asulam was effective in eradicating bracken (<1% cover remaining) on a third of sites. However, on 10% of the sites, the bracken had regenerated completely (cover >80%) and on the remainder it was still present in patches (>20% of quadrats), often at high density. More than half the sprayed sites had seen good recovery of moorland vegetation, the target of the restoration, because they were now classified as having upland heathland vegetation within the National Vegetation Classification. Considerable amounts of bracken control are grant aided as part of agri‐environment schemes. These schemes should be adapted to encourage good practice, namely, intensive follow‐up treatment by spraying any emerging fronds, and to encourage treatment of previously sprayed areas rather than spraying of new areas in order to protect previous investment of grant aid.     

The effects of cutting and herbicide treatment on Pteridium aquilinum encroachment

Abstract. Pteridium aquilinum (bracken) encroachment is an important factor in the loss of certain habitats in the United Kingdom. However, no information exists as to whether prevention of encroachment is a cost‐effective strategy for Pteridium management. Conventional methods for the control of Pteridium (cutting, asulam application) were tested at one site (Levisham) to quantify their ability to prevent or delay encroachment and to affect the vigour of the Pteridium at the edge of the stand. The effects of encroachment and asulam application on the vegetation present were monitored at a second site (Ramsley), where techniques commonly used for moorland restoration were employed in combination with asulam application. Cutting once per year or a single application of asulam delayed the advance of the Pteridium front. At Levisham, the untreated front advanced 2.7 m in 5 yr, while in the same period the cut front advanced 0.88 m and the sprayed front was 1.5 m behind its initial position. At Ramsley, the untreated front invaded 1.8 m in 5 yr, and the sprayed front was again 1.5 m behind its starting position. Both spraying and cutting reduced frond biomass, frond cover and rhizome biomass. Herbicide spraying prevented the loss of Calluna vulgaris, though the restoration treatments had little effect. The merits of a balanced targeting of control on encroaching fronts or Pteridium at the stand level are discussed for different situations.     

The Control of Bracken

Abstract

Bracken is a weed of economic importance to hill farming, conservation, and forestry in many parts of the world. It is an aggressive competitor, capable of rapid spread by means of its persistent underground rhizomatous system, and it is highly toxic and carcinogenic to domestic stock. In addition, it harbors large populations of ticks that transmit disease to animals and humans.

Control by husbandry or by mechanical frond destruction is labor-intensive and often inadequate. Chemical control continues to be a subject of intensive research, with particular emphasis on the use of adjuvants, including surfactants. There is evidence that formulation of systemic herbicides such as asulam or glyphosate with low concentrations (e.g., 0.05%) of certain nonionic surfactants can effectively increase foliar absorption and translocation to the underground rhizome buds.

Biological methods of control using either insects or fungi are under investigation. Certain bracken-feeding South African insects are specific to the variety of bracken (Pteridium aquilinum ssp. aquilinum var. aquilinum) that is also common in the U.K. It is proposed that the moth Conservula consigna would occupy a vacant ecological niche if introduced into the U.K. bracken ecosystem and thereby give effective control. The insect feeds on the pinnae and completes its life cycle on bracken. The fungus Ascochyta pteridis is indigenous to the U.K., where, under certain climatic conditions, it causes the disease of bracken known as curl-tip. The potential of a mycoherbicidal formulation, in which the spores are mixed with adjuvants designed to give the fungus a competitive advantage, is under investigation. Oil and water emulsions, for example, are expected to be of particular importance: they assist adhesion of spores to the host surface while providing water for their germination.

This review examines the problems posed by bracken and the progress of research toward the development of economically and environmentally acceptable methods of control of this noxious and aggressive weed.     

Long-term recovery of bracken (Pteridium aquilinum (L.) Kuhn) after asulam spraying

This paper investigates the effects of a single asulam application, sprayed from the air, on the rhizome biomass, bud density, fronds and carbohydrate reserves of bracken (Pteridium aquilinum (L.) Kuhn) using a time sequence approach. Regression models were used to investigate how these characteristics varied with time after spraying, and were used, where appropriate, to calculate the time taken for full recovery after treatment. Frond density and biomass recovered in approximately eight years, bud numbers in seven, but rhizome biomass and total carbohydrate reserves required 10 to 12 years to recover. The consequences of these results are compared with predictions from a computer model and discussed in relation to the best timing of re-treatment and the management needed for long term control.     

Bracken thiaminase-mediated neurotoxic syndromes

The toxicity of bracken ( Pteridium aquilinum (L.) Kuhn) to animals is complicated because this plant elaborates more than one type of agent harmful to livestock. An enzyme, thiaminase I, which destroys thiamine, is responsible for the neurotoxic syndrome. Using a radiochemical assay, the distribution of thiaminase I activity in bracken throughout the growing season has been ascertained: levels are high in the rhizome and young buds, but fall sharply in the fronds as the aerial parts of the plant unfold.
The so-called thermostable 'antithiamine' factors present in bracken and other plant species are discussed.
The biochemical lesions of thiamine deficiency in animals are briefly outlined, and the clinical syndrome caused by the inclusion of bracken fronds or rhizomes in the diet for simple-stomached animals (rat, horse, pig) and a ruminant (sheep) are described.
All these neurotoxic syndromes respond to thiamine therapy in a dramatic way, if administered during the early stages of the disease.     

The ecological status of bracken

Information about the morphology of bracken ( Pteridium aquilinum (L.) Kuhn var. aquilinum ) and the chief nutrients in the frond at different times of the year introduce an account of litter production and its accumulation in relation to the behaviour of frond, root and rhizome systems.
Where litter gain exceeds loss there is a correlation between the thickness and/or kind of litter and the level of the root and rhizome systems in relation to the mineral soil surface: with increase of litter the bracken becomes progressively more dependent for its physical and chemical soil environment on its own débris and less on the underlying mineral soil. An example of the limit of complete dependence has not been examined, but degeneration of the community can take place before that stage is reached.
From a review of the chief factors affecting bracken the conclusion is reached that the woodland habitat is both favourable and restrictive: in it bracken is in equilibrium with its environment, at a high social status. The relationship with other plant communities depends largely on the degree of human interference to which each is subject. Dominant bracken when left alone, and where gain of litter exceeds loss, becomes the victim of its own success; local degeneration opens the way for entry by other species.     

Translocation of C in Macrocystis pyrifera (Giant Kelp)

The pattern of import and export of ¹⁴C-labeled assimilates in Macrocystis pyrifera (L.) C. A. Agardh in southern California was studied by labeling single blades on fronds, in situ, with [¹⁴C]NaHCO₃ for 24 hours. The pattern was found to be similar to that known in dicotyledons: actively growing tissue imported and did not export. As a blade reached maturity it began to export, at first only acropetally to the apex which formed it, later also down the frond to sporophylls and frond initials at the base of the frond, and into the apical regions of juvenile fronds; finally there was a phase of declining export, late in the life of the blade, when transport was only downward. Young fronds imported from older fronds until they were approximately 3 meters long, by which time they had developed mature, upward exporting blades. No translocation was found from a younger frond to an older frond, nor was there transport upward from a blade on a frond lacking the apical region.     

根茎半灌木羊柴对光合同化物的克隆整合

羊柴为根茎半灌木,主要分布于我国北方半干旱区的沙地。该种的营养繁殖是由根茎的水平延伸使顶端分生组织与腋生分生组织序列化不断产生而进行的。本项研究采用^14C标记和去叶实验,探讨了母株、子株所组成的分株对内的光合同化物的整合。在24h的传输中,相当可观量的14C光合同化物自母株传输给子株、子株的根系和根茎,且它们的汇活度均超过了10%。在^14C-光合化物向基向传输（由子株向母株）中,传输率较小,且汇活度不足1%,但根茎在这一传输中的汇活度却超过2%。去叶影响着母株与子株之间的光合同化物整合。保持完整的母株增大向遭受去叶的子株传输光合产物;反之亦然。母相提并论和子株都向根茎传输一定量的同化物,表明根茎自分株获得光合同化物的支持。羊柴的相连分株间通过根茎而发生克隆整合,可能使其分株和基株在不利的沙地生境中都获得净收益。     

Rhizome phyllosphere oxygenation in Phragmites and other species in relation to redox potential, convective gas flow, submergence and aeration pathways

Underground rhizomes of emergent aquatic macrophytes are important for perennation, vegetative spread, competition and anchorage. In four species we examined the potential for the development of oxidized phyllospheres around rhizome apical buds, similar to the protective oxygenated rhizospheres around roots. Redox potentials and polarographic measurements of radial oxygen loss were recorded using platinum cathodes around the apical buds. The aeration pathway from atmosphere to phyllosphere was investigated anatomically and by applied pressurized gas flow. Redox potentials increased by +400, +45, +200 and +340 mV around rhizome apices of Phragmites australis, Oryza rhizomatis, Carex rostrata and Glyceria maxima, respectively. Radial oxygen loss from rhizome apices of Phragmites was increased by convective gas flow through the rhizome and by shoot de-submergence, and decreased by resistances applied within the aeration pathway and by shoot submergence. We conclude that oxygen passes via internal gas-space connections between aerial shoot, rhizome and underground buds and into the phyllosphere regions via scale-leaf stomata and surfaces on the buds. We suggest that oxidized phyllospheres may protect rhizome apices against phytotoxins in waterlogged soils, just as oxidized rhizospheres protect roots.