Title:
|
Factors contributing to the regeneration of bracken
|
Bracken (Pteridium aquilinum (L. ) Kuhn) is considered a problem plant in many
countries, so an understanding of its method of reproduction is important to those
trying to control it, prevent its spread or to manage rangelands sustainably.
It is commonly believed that in the UK, bracken mainly reproduces asexually via
its rhizome system. However, spores are produced in many locations in some
years and in some places apparently every year. What stimulates the production
of these spores and whether they contribute to the spread of the plant is
unknown. The spores as allergens are hazardous to the health of animals
including humans, and may be carcinogenic when ingested in sufficient dosage.
The aims of this study were therefore to identify both the conditions conducive to
spore production, and the conditions that may promote the growth and survival
of gametophytes and sporelings.
To test what conditions were associated with spore production, three
experiments using bracken sporophytes of different genotypes grown in pots in
controlled environment growth chambers were carried out. The experiments
differed in their designs and the range of variables tested, but all were
multifactorial. The environmental variables were: two different levels of
photosynthetically active radiation (PAR) and light quality (simulated broad-leaf
canopy shade versus simulated full sun), two temperature regimes (each
resembling one of those at the sites inhabited by the source plants), two nutrient
addition rates, two water addition treatments, and rhizome disturbance. A novel
quantitative method of scoring sporangial development was developed and used
to assess the effects of these factors. Genotypeh ad a significant effect on
sporangial development, but not on the rate of that development. Nutrient and water additions affected vegetative growth, but had no significant effects on
sporangial development. The higher of the two rates of PAR and temperature
increased both the rate and the maximum sporangial development attained.
Rhizomes that were excavated and cut to length immediately before the
experiment produced fronds with sporangia that developed at a slower rate and
attained a lower degree of maturity than those rhizomes that had been excavated
and cut prior to the previous growing season. There was no correlation of
biomass of the sporophyte with sporangial development, or development of
sporangial tissue on fronds from rhizomes in a second experiment, with
sporangial development of the same rhizomes in the previous year. This shows
that environmental factors influence sporangial development in bracken, with
genotype having an influence on maximum spore output, but that the rate of
development depends entirely on some environmental variables.
Spores of bracken were cultured in a wide range of different conditions.
The variables included temperature, PAR; state of media; calcium content; pH;
soil water content, and relative humidity. Of these, only temperature
significantly affected germination, with 20°C producing the highest germination
of 5,10,15,18,20,25 and 30°C. Gametophytes were more likely to reach a state
of 2-dimensional growth (transition) at 20 or 25°C than at 18 or 30°C. Lower
levels of PAR decreased the likelihood of gametophytes reaching transition. Of
the calcium levels tested, only complete lack of added calcium had a significant
effect on gametophytes, with increased gametophyte mortality and a decrease in
numbers reaching transition. Media at pH 4 and 8 reduced numbers of
gametophytes reaching transition compared to pH 6.25. There was increased
mortality at pH3 Water content of the substratum outwith the range 50 - 150% field
capacity, significantly reduced the number of cordate gametophytes. Humidities
in the range 10 - 100% relative humidity did not have any significant effect on
gametophyte growth. This may have been due to the constant availability of
water from the substratum. Conditions conducive to gametophyte development to
transition were; a temperature above 8°C, the presence of calcium, the absence of
deep shade, a pH near to 6.25 and adequate soil moisture.
Data from the growth chamber and spore culture experiments involving
temperature were used to create a model of bracken sporulation and spread in the
British Isles. Resultant maps show that the south east is at the highest risk of
bracken spread by spores, however, when climate change predictions are used in
conjunction with the model, it can be seen that northerly areas may be at risk in
the future.
This research shows that while genotype does play a part in determining
spore production of bracken, environment plays the bigger role. It is likely that
with the predicted increase in temperatures in the UK, more bracken plants will
begin to spore in more areas of the country, putting more animals and people at
risk from health problems and potentially further degrading susceptible land.
This work does not explain why bracken gametophytes are rarely encountered in
field situations. Since the conditions that were found to be conducive to
gametophyte establishment are often found in the UK, either one or more factors
that were not included in these experiments play a key role in determining the
success of sporal regeneration, or colonisation by spores occurs more frequently
than is widely believed.
|