A comparison of the growth curves of the foliose lichen Parmelia conspersa determined by a cross-sectional study and by direct measurement

Changes in the radial growth rate (RGR mm/yr) through life were studied in thalli of the foliose lichen Parmelia conspersa by two methods: (1) a cross-sectional study (Study A) in which the RGR was measured in 60 thalli from 0.2 to 13 cm in diameter, and (2) by radial growth measurements over 4.5 years of fragments, consisting of a single major lobe, which were removed from large thalli and glued to pieces of slate (Study B). Both studies suggested there was a phase of increasing RGR in small thalli followed by a more constant phase, the latter beginning at approximately a thallus radius of 6-8 mm. However, in Study B significantly increased RGR was observed during the second 6-month growth period. This phase of growth was more likely to be due to an increase in lobe width than to an effect of climate. In addition, a lobe in a large thallus with both adjacent lobes removed significantly increased in width over 1 year compared with control lobes. These results suggest that (1) mean lobe width in a thallus may be determined by the intensity of marginal competition between adjacent lobes, and (2) changes in lobe width during the life of a lichen thallus may be a factor determining the establishment of the linear phase of growth in foliose lichens. © 1992.

The response of lichen growth to additions of distilled water, rainwater and water from a rock surface

Three lichen species were wetted in the field with distilled water, rainwater or water which had run off a rock surafce, during July 1974 to February 1975. The radial growth rate of Parmelia glabratula ssp. fuliginosa was not influenced by the wetting treatments. The radial growth rate of P. conspersa with the distilled water was greater than the control, rainwater and runoff treatments. The radial growth rate of Physcia orbicularis was lower with rainwater and runoff treatmentss than the control and distilled water treatment. These results may be explained by the effect of wetting on the carbon balance of the lichens and by the influence of water chemistry.

The reponse of lichen growth to transplantation to rock surfaces of different aspect

Thalli of four saxicolous lichens on slate rock fragments were transplanted from rock surafces to horizontal boards and then to south-east-facing and north-west-facing rock surfaces. The radial growth rate of Physcia orbicularis and Parmelia conspersa thalli declined after transplatation to north-west-facing rock surfaces and was unchanged after transplantation to south-east-facing rock surfaces cmpared with growth rates on the boards. The radial growth rate of P. glabratula ssp. fuliginosa thalli declined after transplantation to south-east-facing rock surfaces and was unchanged after transplantation to north-west-facing rock surfaces compared with grwoth rate on the boards. The radial growth rate of P. saxatilis thalli was similar on the horizontal boards, south-east-facing and north-west-facing rock surfaces. These results are dsicussed in relation to the aspect distribution of the four lichens in South Gwynedd, Wales.

The influence of the frequency of wetting and drying on the radial growth of three saxicolous lichens in the field

Three lichen species were wetted with distilled water at different frequencies during August 1973 to July 1974. The radial growth rates of Parmelia glabratula ssp. fuliginosa and Physcia orbicularis thalli declined with increased wetting while the radial growth rate of Parmelia conspersa thalli increased with wetting frequency until ten experimental wettings per month but at fifteen wettings per month fell to a value near to the control. In the summer months, wetting resulted in a decline in the radial growth of P. glabratula ssp fuliginosa compared with the control but had little influence on the growth of P. conspersa and Physcia orbicularis. In the winter months, wetting had no significant influence on the radial growth of Parmelia glabratula ssp. fuliginosa, while the radial growth of P. conspersa increased and Physcia orbicularis declined compared with controls. These results are interpreted physiologically and in relation to the aspect distribution of the three lichens on rock surfaces.

Studies on the growth rates of lichens

Progress in the field of lichen growth rates is briefly reviewed. The application of a new method of measuring growth rate to thalli of different size has led to the conclusion that there are changes in the radial growth rate during the life of a lichen thallus. For most of the life of a lichen thallus the radial growth rate is constant and the thallus radius increases linearly. Preceeding the linear phase the radial growth rate increases with time and the thallus radius increases logarithmically. There is no evidence for a postlinear phase in the radial growth of a lichen thallus. Studies on the growth rate of lichens are applied both to the problems of determining the age of a lichen thallus on an undated substratum and to an ecological investigation in the field.

The influence of aspect on the pattern of saesonal growth in the lichen Parmelia glabratula ssp. fuliginosa (Fr. ex Duby) Laund

The radial growth of samples of thalli of Parmelia glabratula ssp. fuliginosa were measured in situ on a south-facing and a northwest-facing rock surface each month from August 1973 to July 1974. In the periods August to October 1973 and March to July 1974 the radial growth of thalli in the northwest population was greater than in the south population. In the period November 1973 to February 1974 the radial growth of thalli in the south population was greater than in the northwest population. A physiological basis for the differences in seasonal growth in the two populations was suggested. The mean annual radial growth rate (in units of mm/year) was not significantly different in the two populations. However, the variability in radial growth rate between thalli was signifiacntly larger in the northwest than in the south population. These results may be explained by genetic difference between the populations and environmental differences between the rock surfaces.

Studies on the growth rates of lichens

Progress in the field of lichen growth rate studies is briefly reviewed. The application of a new method of measuring growth rate to thalli of different size has led to the conclusion that there are changes in the radial growth rate during the life of a lichen thallus. For most of the life of a lichen thallus the radial growth rate is constant and the thallus radius increases linearly. Preceding the linear phase the radial growth rate increases with time and the thallus radius increases logarithmically. There is no evidence for a postlinear phase in the radial growth of a lichen thallus. Studies on the growth rate of lichens are applied both to the problems of determining the age of a lichen thallus on an undated susbtratum and to an ecological investigation in the field.

Chemical studies of plasma etchants used in integrated circuit manufacture

Plasma or "dry" etching is an essential process for the production of modern microelectronic circuits. However, despite intensive research, many aspects of the etch process are not fully understood. The results of studies of the plasma etching of Si and Si02 in fluorine-containing discharges, and the complementary technique of plasma polymerisation are presented in this thesis. Optical emission spectroscopy with argon actinometry was used as the principle plasma diagnostic. Statistical experimental design was used to model and compare Si and Si02 etch rates in CF4 and SF6 discharges as a function of flow, pressure and power. Etch mechanisms m both systems, including the potential reduction of Si etch rates in CF4 due to fluorocarbon polymer formation, are discussed. Si etch rates in CF4 /SF6 mixtures were successfully accounted for by the models produced. Si etch rates in CF4/C2F6 and CHF3 as a function of the addition of oxygen-containing additives (02, N20 and CO2) are shown to be consistent with a simple competition between F, 0 and CFx species for Si surface sites. For the range of conditions studied, Si02 etch rates were not dependent on F-atom concentration, but the presence of fluorine was essential in order to achieve significant etch rates. The influence of a wide range of electrode materials on the etch rate of Si and Si02 in CF4 and CF4 /02 plasmas was studied. It was found that the Si etch rate in a CF4 plasma was considerably enhanced, relative to an anodised aluminium electrode, in the presence of soda glass or sodium or potassium "doped" quartz. The effect was even more pronounced in a CF4 /02 discharge. In the latter system lead and copper electrodes also enhanced the Si etch rate. These results could not be accounted for by a corresponding rise in atomic fluorine concentration. Three possible etch enhancement mechanisms are discussed. Fluorocarbon polymer deposition was studied, both because of its relevance to etch mechanisms and its intrinsic interest, as a function of fluorocarbon source gas (CF4, C2F6, C3F8 and CHF3), process time, RF power and percentage hydrogen addition. Gas phase concentrations of F, H and CF2 were measured by optical emission spectroscopy, and the resultant polymer structure determined by X-ray photoelectron spectroscopy and infrared spectroscopy. Thermal and electrical properties were measured also. Hydrogen additions are shown to have a dominant role in determining deposition rate and polymer composition. A qualitative description of the polymer growth mechanism is presented which accounts for both changes in growth rate and structure, and leads to an empirical deposition rate model.

The use of lichen growth rings in lichenometry: some preliminary findings

Certain species of crustose lichens have concentrically zoned margins which probably represent yearly growth rings. These marginal growth rings offer an alternative method of studying annual growth fluctuations, establishing growth rate-size curves, and determining the age of thalli for certain crustose species. Hence, marginal growth rings represent a potentially valuable, unexploited, tool in lichenometry. In a preliminary study, we measured the widths of the successive marginal rings in 25 thalli of Ochrolechia parella (L.) Massal., growing at a maritime site in north Wales. Mean ring widths of all thalli varied from a minimum of 1.02 mm (the outermost ring) to a maximum of 2.06 mm (the third ring from the margin). There is some suggestion that marginal ring width and thallus size are positively correlated; and hence that growth rates increase in larger thalli in this small population. In a further study on recently exposed bedrock adjacent to Breidalon, SE Iceland, we examined the potential for using marginal growth rings to estimate thallus age of a lichen tentatively identified as a Rhizocarpon (possibly R. concentricum (Davies) Beltram.) and thus confirm the timing of surface exposure (c. 50 years). Collectively, these results suggest: 1) the measurement of marginal rings is a possible alternative method of studying the growth of crustose lichens; 2) O. parella may grow differently to other crustose species, exhibiting a rapidly increasing radial growth rate in thalli >40 mm; 3) where lichens with marginal rings grow on recently exposed surfaces (<60 yrs), minimum age estimates can be made using growth rings as an in situ indication of lichen growth rate; 4) it is suggested that this phenomenon could provide a valuable, previously unexploited, in situ lichenometric-dating tool in areas lacking calibration control.

Growth of crustose lichens: a review

Crustose species are the slowest growing of all lichens. Their slow growth and longevity, especially of the yellow-green Rhizocarpon group, has made them important for surface-exposure dating (‘lichenometry’). This review considers various aspects of the growth of crustose lichens revealed by direct measurement including: 1) early growth and development, 2) radial growth rates (RGR, mm yr-1), 3) the growth rate-size curve, and 4) the influence of environmental factors. Many crustose species comprise discrete areolae that contain the algal partner growing on the surface of a non-lichenised fungal hypothallus. Recent data suggest that ‘primary’ areolae may develop from free-living algal cells on the substratum while ‘secondary’ areolae develop from zoospores produced within the thallus. In more extreme environments, the RGR of crustose species may be exceptionally slow but considerably faster rates of growth have been recorded under more favourable conditions. The growth curves of crustose lichens with a marginal hypothallus may differ from the ‘asymptotic’ type of curve recorded in foliose and placodioid species and the latter are characterized by a phase of increasing RGR to a maximum and may be followed by a phase of decreasing growth. The decline in RGR in larger thalli may be attributable to a reduction in the efficiency of translocation of carbohydrate to the thallus margin or to an increased allocation of carbon to support mature ‘reproductive’ areolae. Crustose species have a low RGR accompanied by a low demand for nutrients and an increased allocation of carbon for stress resistance; therefore enabling colonization of more extreme environments.

Factors affecting the growth of Cladophora in relation to river pollution

Nuisance growths of Cladophora have been associated with eutrophication. A review of the literature, however, reveals a scarcity of relevant experimental growth studies. Sampling experimental streams reveals that the addition of sewage effluent to good quality water alters the flora from that dominated by Potamogetan crispus to one dominated by CLadophora. Spatial and temporal differences in biomass of taxa present are discussed in the context of accompanying physicochemical data. In laboratory batch culture, growth of unialgal C. glomerata was accompanied by elevation of medium pH - considered largely responsible for the poor growth in such culture. However, appropriate experimental conditions and indices of growth were selected and the effects of various herbicides assessed. Diquat and terbutryne were shown to possess algicidal activity towards Cladophora. A closed continuous culture apparatus was developed: growth proceeded through lag, logarithmic and linear phases. Inoculum size and medium flow rate had significant effects on growth, and were standardized. In continuous culture, specific growth rate increased linearly with increased duration of light per day, up to 24 hours, and increased light intensity, up to 6000 lux - the highest intensity tested. Comparison of field and laboratory results suggests that ammonia toxicity is attributable to the undissociated form. In the laboratory, 185 µg/1 undissociated ammoniacal nitrogen reduced specific growth rate to 50% of that at 10 µg/1 undissociated ammcniacal nitrogen. 0.077-1.057 mg/1 NO2-N had no significant effect on growth. 7.2-15.2 mg/1 NO3-N had no significant effect on specific growth rate. Neither was any nitrate/phosphate interaction significant. At 4.9 mg/1 PO4-1, specific growth rate was only 48% of that at 1.9 g/1 P04-P. The critical medium PO4-P concentration was <0.1 mg/i. Specific growth rate was reduced to 50% of that in natural water by 0.036 mgCu/l, 0.070 mgzn/1 and 1.03 mgPb/l. Metal uptake was evaluated.

Studies on biofilm growth of Pseudomonas aeruginosa PAO1

The development of in vitro techniques to model the surface-associated mode of growth is a prerequisite to understanding more fully the physiological changes involved in such a growth strategy. Key factors believed to influence bacterial persistence in chronic infections are those of the biofilm mode of growth and slow growth rate. Methods for controlling Pseudomonas aeruginosa biofilm population growth rates were investigated in this project. This microorganism was incompatible with the in vitro 47mm diameter membrane filter-based biofilm technique developed for the study of Escherichia coli and Staphylococcus epidermidis by Gilbert et al (Appl. Environ. Microbiol. 1989, 55, 1308-1311). Two alternative methods were designed. The first comprised a 25mm diameter cellulose acetate membrane filter supported in an integral holder. This was found to be limited to the study of low microbial population densities with low flow rates. The second, based on a cylindrical cellulose fibre depth filter, permitted rapid flow rates to be studied and allowed growth rate control of biofilm and eluted cells. Model biofilms released cells to the perfusing medium as they grew and divided. The viability of released cells was reduced during, and shortly after, inclusion of ciprofloxacin in the perfusate. Outer membrane profiles of biofilm populations exhibited at least two bands not apparent in planktonic cells grown in batch and chemostat culture, and LPS profiles of biofilm populations showed variation with growth rate. Cell surface hydrophobicity of resuspended biofilm cells varied little with growth rate, whilst it decreased markedly for cells released from the biofilms as growth rate increased. Cells released from the biofilm were more hydrophilic than their sessile counterparts. Differing growth rates, LPS profiles and hydrophobicity are proposed to have a bearing on the release of cells from the adherent population.

Creep crack growth in cast steam turbine casing steels

The creep rupture properties of cast ½Cr½Mo¼V and 1Cr1Mo¼V alloy steel used in the manufacture of power station steam generating plant. have been investigated. The effects of constraint and geometry on the creep rupture properties are also considered. The validity of various criteria controlling macroscopic creep crack growth in cast CrMoV alloys has been examined. It is found that neither the stress intensity factor nor reference stress correlate satisfactorily the creep crack growth rates at the test temperature of 550°C. Certain minimum displacements must be achieved for crack initiation and propagation. It is found that this displacement as measured by crack opening displacement or crack aspect ratio, is the same in both compact tension and centre-cracked panel geometries, is invariant with crack length and decreases with increasing constraint. The effect of constraint on creep crack growth rate in the two geometries is less conclusive. A new model describing creep crack growth in cast CrMoV alloy steels has been developed. The model is based on the results from a numerical finite element creep analysis of the relaxation and redistribution of stress ahead of an incubating creep crack . It is found that macroscopic creep crack growth in a material undergoing either plane stress or plane strain deformation can be described by a fracture stress which is based on the Von Mises equivalent stress. It has been shown that this model is capable of rationalising all of the experimental crack velocity data from the cast CrMoV alloys. The resultant degree of data correlation is far superior to that obtained when using the stress intensity factor or reference stress. A cumulative damage creep fracture model based upon the results from the numerical analysis has been developed. It is found that the model is capable of predicting the behaviour of propagating creep cracks in cast CrMoV alloys from smooth bar creep rupture data.

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr-1. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.

The biology of the lichen Rhizocarpon geographicum: symbionts, growth, ecology, and lichenometry

Rhizocarpon geographicum (L.) DC. is one of the most widely distributed species of crustose lichens. This unusual organism comprises yellow-green ‘areolae’ that contain the algal symbiont which develop and grow on the surface of a non-lichenized, fungal ‘hypothallus’ that extends beyond the margin of the areolae to form a marginal ring. This species grows exceptionally slowly with annual radial growth rates (RGR) as low as 0.07 mm yr-1 and its considerable longevity has been exploited by geologists in the development of methods of dating the age of exposure of rock surfaces and glacial moraines (‘lichenometry’). Recent research has established some aspects of the basic biology of this important and interesting organism. This chapter describes the general structure of R. geographicum, how the areolae and hypothallus develop, why the lichen grows so slowly, the growth rate-size curve, and some aspects of the ecology of R. geographicum including whether the lichen can inhibit the growth of its neighbours by chemical means (‘allelopathy’). Finally, the importance of R. geographicum in direct and indirect lichenometry is reviewed.