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.     

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 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 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.     

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 architecture of a dimorphic clonal fern, Lomagramma guianensis (Aublet) Ching (Dryopteridaceae)

Lomagramma guianensis possesses a dimorphic rhizome and trimorphic fronds. The juvenile genet consists of a slender, sinuous, unbranched, terrestrial rhizome which bears pinnatifid fronds. The adult rhizome form is initiated when the apex becomes scandent. It is stout, exhibiting rhythmic production of pinnate, adult-type fronds, and probably rhythmic growth. The scandent rhizome is ultimately monocarpic, producing fertile acrostichoid fronds and subsequendy dying. Terrestrial runners are produced from proleptic buds at the scandent rhizome base. These are largely similar in morphology to the scandent rhizomes, but display different behaviour. The runners are relatively ephemeral structures which produce no fronds. They branch proleptically from their base to produce scandent rhizomes, and also along their length to give rise to a complex system of runnerS. Senescent runners which have lost contact with their parent scandent rhizome produce reiterative complexes which represent a return to the juvenile form. These reiterated rhizomes behave identically to the juvenile genets, and repeat the entire cycle of development. The architectural history of the clonal pteridophyte Lomagramma is compared to that of angiosperm trees; interesting similarities and differences are found.     

The mode of action of asulam [methy] (4-aminobenzenesulphonyl) carbamate] in bracken

Foliage-application of 4.4 kg/ha asulam [methyl (4-aminobenzenesulphonyl) carbamate] to field bracken reduced frond density by 97.1, 87.6 and 77.8%, one, two and three years respectively, after treatment. Examination of the rhizome system from these plots revealed that a high proportion of the rhizome apices and frond buds were killed, the remainder being damaged or remaining dormant. Autoradiography of bracken plants grown from sporelings or fragments and treated with 14 C-ring-labelled asulam revealed that uptake was rapid and progressive with time, whilst translocation was generally basipetal showing a typical source-sink pattern, with intense accumulation in the apices and buds. Optimum penetration and basipetal translocation was achieved when treatments were applied to almost fully-expanded fronds; application to immature fronds resulted in predominantly acropetal movement. Basipetal translocation was extensive in field bracken, accumulation occurring in the buds distant from the base of the treated fronds. There was evidence that asulam was relatively persistent in the rhizome system, 59 and 8% of the applied activity remaining in the rhizome, one and ten months respectively, after treatment. Hydrolysis of asulam produced small amounts of sulphanilamide (<10%) and traces of sulphanilic acid, 4-aminophenol and benzene sulphonamide. Studies on the effect of asulam on the incorporation of 14 C-labelled precursors into RNA and protein indicate that inhibition of RNA and protein syntheses may be major sites of action of asulam.     

扎龙湿地芦苇种群不同龄级根茎构件的季节动态

克隆植物根茎具有营养繁殖和扩展种群的功能,也是芽和分株生理整合的通道。根茎构件具有出生、死亡及年龄等种群统计特征,不同龄级根茎的季节动态可以反映根茎的存活和衰老过程。采用单位土体挖掘取样,对扎龙湿地4个生境芦苇种群根茎构件进行野外调查,比较不同龄级根茎长度、生物量和干物质贮量的季节动态。结果表明:7—10月份,1a根茎长度、生物量和干物质贮量均呈指数函数增加,在生长季中后期有一个持续时间较长的生长和物质积累时期。6—10月份,2a、3a根茎长度呈线性函数增加,4—6a根茎长度呈线性函数减少;2—4a根茎生物量和2—5a根茎干物质贮量呈二次函数先减少后增加,5a、6a根茎生物量和6a根茎干物质贮量呈幂函数减少。整个生长期内,根茎长度和根茎生物量均以3a最大,根茎长度以最高的6a最小,根茎生物量以最低的1a最小;根茎干物质储量以5a最大,以最低的1a最小。4个生境芦苇种群根茎长度、生物量和干物质贮量在龄级间的差异及差异序位稳定,在新根茎的产生、老根茎的存活以及根茎寿命与养分消耗和储藏上均具有稳定的生物学特性,不同龄级根茎在种群中的地位和作用以及对种群的贡献不同。     

Effect of herbicides on bracken rhizome survival

Glyphosate sprayed at 1.1, 2.2, 4.5 or 6.7 kg/ha on to bracken decreased the total carbohydrate content of the storage rhizome 20 months after spraying, the decrease being greater when spraying was carried out at the beginning rather than at the end of August. Asulam, benazolin and metribuzin, although reducing frond numbers, showed no significant effect on the rhizomes over the same period.     

Distribution of glycolipids and phospholipids in Pteridium aquilinum

The glycolipids and phospholipids in fronds and rhizomes of Pteridium aquilinum were determined. The total quantity of polar lipid decreased towards the base of the frond, but increased in the storage rhizome. The monogalactosyl diglyceride/digalactosyl diglyceride ratio was 1.8 in the pinnae, 1.0 in the lower petiole and 0.3 in the storage rhizome.     

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.     

扎龙湿地不同生境芦苇种群根茎数量特征及动态

采用单位土体取样,计测长度和生物量的调查与统计方法,对扎龙湿地保护区4个生境芦苇种群根茎数量特征进行比较分析。结果表明,芦苇5月10日左右返青后进入营养生长期,根茎长度6—8月份缓慢增加,8—10月份显著增加,后期是前期的3.5—10.3倍,生长季中后期是种群新根茎补充和生长的主要时期,不仅实现了种群空间扩展,并为营养繁殖储备更多繁殖芽;根茎生物量和干物质贮量6—8月份逐渐减少,8—10月份又逐渐增加,均以生长季末期的10月份最大,并均显著地(P0.05)高于其他月份,种群根茎养分的消耗主要供给根茎芽的萌发和幼株生长,根茎养分的储藏又为翌年种群的更新及扩展提供物质保障,种群对地下根茎存在明显的养分"超补偿性"贮藏现象。种群根茎长度和生物量均以湿生生境最大,依次为旱生生境、水生生境,盐碱生境最小,根茎干物质贮量以旱生生境最大,依次为湿生生境、水生生境,盐碱生境最小。种群根茎长度与返青后实际生长时间之间均较好地符合直线函数关系,种群根茎生物量和干物质贮量与生长时间之间较好地符合二次曲线函数关系,R2在0.804—0.997之间,拟合方程均达到了显著或极显著(P0.01)水平。4个生境芦苇种群在根茎长度、生物量、干物质贮量等数量特征均表现出由遗传因素控制的比较稳定的季节动态规律,在生境间的差异及其差异序位又均基本稳定,均表现出明显的土壤因子环境效应,其中土壤含水量、有机质、速效氮为正向驱动,p H、速效磷为负向驱动,土壤含水量、p H对根茎数量特征的驱动作用更突出。     

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.     

Aquilinan,an acidic polysaccharide from Pteridium aquilinum

A water-soluble, acidic polysaccharide, aquilinan, was isolated from storage rhizomes and young fronds of bracken and shown to be homogeneous. The polysaccharide contained galactose, xylose, fucose and arabinose residues together with chains of repeating α(1-2′)glucuronosylmannose units. A method of correcting the mannose content for degradation during the hydrolysis of the aldobiuronic acid is described.     

Seasonal changes in the amount of carbohydrate and crude protein in the rhizome ofMiscanthus sacchariflorus

In the herbaceous perennials, seasonal changes in the amount of reserve substances play an important role in the production processes of whole plants. Mutohet al. (1968) indirectly estimated the amount of reserve substances stored in the rhizome ofMiscanthus sacchariflorus by measuring the bulk density of the rhizome, and found aarked seasonal changes. In the present study seasonal changes in the carbohydrate and crude protein content of the rhizome were determined and the results obtained were compared with the changes in the total amount of reserve substances estimated by means of changes in the bulk density. A good parallel seasonal relationship was confirmed between the amount of consumable carbohydrate determined chemically and the quantity of reserve substances estimated from the changes in bulk density. The total yields of carbohydrate and crude protein were ca. 75% of the available reserve substances estimated from the changes of bulk density for the old rhizomes and ca. 65% for the new rhizomes in the middle of December.     

CHANGES IN STARCH DISTRIBUTION IN THE OVERWINTERING ORGANS OF TYPHA LATIFOLIA (TYPHACEAE)

Histochemical determinations for storage of carbohydrates in rhizomes, roots, and young shoots of Typha latifolia L. (Typhaceae) were conducted during the overwintering period from November to April. Early winter analysis showed that rhizomes and roots contained large amounts of starch (45.03% and 22.80% dry weight, respectively). The major storage tissue was parenchyma of the rhizome central core. From winter into spring a gradual decrease in storage starch in the rhizome and root occurred concurrently with starch accumulation near zones of rapid development in young shoots (buds), but the rhizome retained much starch (27.40% dry weight) into the start of its 2nd yr.     

松嫩平原碱化草甸旱地生境芦苇种群的芽流和芽库动态

在松嫩平原碱化草甸,旱地生境芦苇种群的根茎分布在土层约1m的不同深度,一般可生活6个年度,个别根茎可存活7～9年,乃至更长的时间.通过芦苇根茎芽调查,创建了植物种群的芽流模型.提出了采用当年1龄级根茎芽的输入率与其它龄级休眠芽库存率之和估计芦苇种群芽库贮量动态的方法.结果表明,随着生长季的进程,芦苇种群芽库输入率呈不断增加趋势,而萌发输出率呈不断减少的趋势,死亡输出率则大体保持相同的较低水平.至休眠前期的9月底,芽库输入率已为输出率的2.04倍.在松嫩平原碱化草甸旱地生境,芦苇种群各龄级根茎的休眠芽有一个稳定的萌发输出过程.定量分析结果表明,芦苇种群不同龄级根茎的休眠芽每年都有11%的比率萌发形成1龄级新根茎.1龄级根茎顶端翌年发育为分蘖株后,可为直接相连接的老龄级根茎就近输送养分,从而实现老龄级根茎芽的活力.     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

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 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.