A STUDY OF GROWTH RINGS IN TREES

The frequency of ‘lines’ of denser wood within theannual ring in larch (Larix decidua Mill.) was compared withmonthly records of rainfall and tables of droughts and rain-spellsin the Bristol region. Moderate correlations were found withJune rainfall (inverse), days in summer droughts, and a ratiogiving a measure of the maximum dry-wet contrast between successivemonths from May to August. This ‘contrast ratio’showed a high degree of parallel variation with ‘line’frequency. A study of individual years suggested that the formationof a pronounced ‘line’ is associated with the occurrenceof a spell of dry weather followed by a wet spell during thelater part of the summer. ‘Mid-lines’ were foundto be associated with a June drought breaking in July; late‘lines’ with a dry-wet contrast in July or August,but in some trees these may be suppressed by the stoppage ofgrowth. Vaguer ‘lines’ of lower frequency were associatedin certain years with wet spells not preceded by noticeabledrought. Individual ‘lines’ can thus be identifiedand roughly dated by some weather feature, so that their positionsheds some light on the length of the growing season. It issuggested that ‘lines’ of denser wood may be occasionedby the killing or inactivating of mycorrhizal rootlets eitherby drought or by waterlogging, followed by partial recoverywhen the weather changes. This would accord with the theorythat summer wood formation is normally associated with a developingwater-deficit in the wood.     

Effect of Late-season Top-dressings of N (and K) applied to Conifer Transplants in the Nursery on their Survival and Growth on British Forest Sites

At two English forest nurseries, transplants of five coniferspecies—Picea sitchensis, Picea abies, Tsuga heterophylla,Abies grandis, and Pinus contorta—were grown with fertilizersupplying N, P, K, and Mg in amounts intended to be adequatefor producing healthy green trees with nutrient concentrationsin the ‘sufficiency range’ as determined by earlierexperiments. ‘Luxury uptake’ of nitrogen was obtainedwith top-dressings of ‘Nitro-Chalk’ applied in thenursery during early September, when top growth had nearly ceased.Tests of the effect of this extra N on forest establishmentwere repeated in four successive years under a wide range ofsoil and climatic conditions, keeping the trees in a cold storeduring each winter and planting them on forest sites in England,Scotland, or Wales during the following spring. Except for Grandfir, nitrogen advanced bud-break of all species during the firstsummer after planting and had no deleterious influence on survival.It tended to increase growth of Sitka spruce during the seasonafter planting, but in later years the differences became smallin relation to tree size. The effects on other species weresmall, except for one considerable decrease in the growth responseof Grand fir at a single site. Frost damage of Sitka spruce of Washington origin was severeon a Welsh and a Scottish site where this frost-sensitive provenancewould not normally be grown. At the Welsh, but not the Scottishsite, the nitrogen treatment increased the damage. In the few experiments (confined to Picea sitchensis) whichtested late-season potassium in the nursery, K concentrationswere increased from deficiency to barely sufficiency level;growth in the forest was increased in two of the four experiments.The extra K had no effect on frost damage.     

Hybrid poplars: present status and potential in Britain

Past and potential future use of hybrid poplars for wood productionin Britain is considered, and preliminary results of trialswith new clones imported from Belgium in 1985 are given. A seriesof nine field experiments established in 1991, to a common protocol,was analysed using stepwise multiple regression. The superiorvigour of ‘Beaupré’ compared with ‘Ghoy’,‘Robusta’ and ‘Trichobel’ was clearin almost every case. For ‘Beaupré’ (Populustrichocarpa x P. deltoides) altitude emerged as the variableexplaining most variation in tree height after six growing seasons,height declining with increasing altitude. The four best siteswere characterized by low elevation and low rainfall. At the two older sites planted in 1987, at Ampthill and Bedgebury,the ‘interamerican’ hybrids (P. trichocarpa x P.deltoides) were the most vigorous, and the clones ‘Unal’and ‘Raspalje’, which were thought to be too rustsusceptible to release for commercial use in 1990, achievedsimilar stem sizes to those of ‘Beaupré’and ‘Bolelare’. The General Yield Class (based ona reference spacing of 8 x 8 m) calculated for ‘Beaupré’was 8 at Ampthill and 24 at Bedgebury. Future prospects for poplar planting are considered in relationto possible reform of the Common Agricultural Policy.     

The Development of Stain in Wounded Sitka Spruce Stems

In order to investigate the severity of staining in woundedSitka spruce stems, the vertical extent of two categories ofstain (based on colour and termed ‘light’ and ‘heavy’)was measured in stems at two sites in south Scotland. Data wereobtained from 98 wounds most of which were between 8 and 14years old and most of which had been caused by Red deer. Theupward extent of stain and its rate of upward spread were positivelycorrelated with wound surface area and length. Most stain wasof the light type which, although it was associated with nearlyall wounds, usually constituted an insignificant defect anddid not commonly extend for more than 1 m ahve wounds. Heavystain occurred above less than half of all wounds studied butwas common above wounds exceeding 300 cm² surface area. Forall wounds, the mean value for the upward extent of heavy stainwas 16.1 cm with a mean rate of spread of 1.9 cm y^–1.However, for wounds exceeding 300 cm² surface area, the valueswere 51.7 cm and 5.5 cm y^–1. These results suggest that,although there is an important effect of wound size on the typeand extent of stain, wounds of the type studied are unlikelyto lead to severe stain and decay in Sitka spruce over a periodof 8–14 years. The results are discussed in relation toother studies on wound staining in conifers.     

Overstorey density influence on the height of Picea abies regeneration in northern Sweden

The study aimed specifically at investigating if canopy opennesswas a better predictor of the height growth of Norway spruce(Picea abies (L.) Karst.) advance regeneration than overstoreybasal area or overstorey standing volume. In 1990, a field experimentwith 3 x 2 factorial design and two replications (blocks) wasestablished in an uneven-aged Norway spruce forest. Plots hada net plot area of 30 x 30 m, each with a 10-m-wide treatedbuffer zone. Three overstorey density levels retained approximately15, 40 and 70 per cent of the pre-harvest overstorey standingvolume and were allotted to the plots. Two types of thinningthat harvested smaller trees or harvested larger trees wererandomly allocated to each pair of overstorey density plots.In mid-June 2000, canopy openness was estimated from hemisphericalphotographs taken at five marked points in the centre of eachof the plots at 0.9 m from ground to the top of the ‘fish-eye’camera lens. Regression results showed that canopy opennesswas a better predictor of height increments of spruce seedlings(0.1< height < 0.5 m), saplings (0.5 height < 2.0m), and small trees (height 2.0 m, diameter at 1.3 m height< 5 cm) than with overstorey basal area (m² ha^–1) oroverstorey standing volume (m³ ha^–1). The height incrementof the spruce advance regeneration was not significantly correlatedto stand basal area or to standing volume. Overstorey basalarea in the net plots was significantly negative (P 0.05) withmean canopy openness estimates, and the r² value was 0.40. Resultsindicated that basal area was not linearly related to canopyopenness as it increased, which might explain the lack of predictivepower of retained basal area on spruce regeneration height indense stands in boreal Sweden.     

CORRIGENDA

Literature References. Wright and Will. Against Scott, R. O.,and Ure, A. M. (1957): Delete ‘Spectrophotometry’and substitute ‘Photometry’. Delete, N.Z.J. Sci.Tech. B. XXXVIII. 699–706 and substitute ‘To appearin The Analyst’.     

The Effect of Drying Quercus robur Acorns to Different Moisture Contents, followed by Storage, either with or without Imbibition

Freshly harvested Quercus robur acorns were spread out, twodeep, in trays, and dried/stored (at +2°C over the courseof 28 weeks) to either 45, 40, 35, 30 or 25 per cent moisturecontent (fresh weight basis). Drying/storage significantly reducedthe germination percentage (P< 0.01). The acorns at eachmoisture content were then either ‘soaked’ or ‘notsoaked’, before immediate germination; or returned tothe same cold store at +2°C, this time in loosely tied polythenebags. Soaking raised the acorns moisture content (M.C.), andalso brought about a significant increase in germination capacity(P<0.01). Regardless of the moisture content immediatelybefore soaking, "soaked" acorns in loosely tied polythene bagsstored better than ‘unsoaked’ acorns. All ‘bagged’acorns (whether ‘soaked’ or ‘unsoaked’)stored better than acorns dried/stored in open trays.     

Some Aspects of the Growth and Spread of Fire in the Open

This paper describes some recent experimental and theoreticalwork on the growth and spread of fire in the open and discussessome examples of field data in terms of the theoretical calculationspresented. The lengths of flames from laboratory fires have been relatedto the size and rate of burning of the fuel by formulae derivedfrom a simplified dimensional analysis. The effects of a windblowing across a long fuel bed on the length and orientationof flames are also described. The scaling laws for flame heightsuggest that in the horizontal spread of fire, heat transferfrom the flames above the fuel bed is important primarily withshallow fuel beds. It is suggested that the main effect of a wind on crib firesis aerodynamic. The wind deflects the advancing fire front fromthe vertical, but perpendicular to this deflected front therate of spread of fire, at least for cribs, is roughly the sameas in still air. However, a theory of spread allowing for heattransfer through the fuel bed and radiation from the flamespredicts that there can be a stable ‘fast‘ spreadas well as a ‘slow’ spread. In ‘fast’spread the flames are thick and control the spread. In ‘slow’ spread radiant heat transfer from theburning zone is usually responsible for the spread. The flamesare thin and of low emissivity. The most important factors determiningthe rate of ‘slow’ spread R are ^pb the bulk densityof the fuel bed and ø the deflection from the verticalof the front of burning fuel which varies with wind speed. Rpb cos ø is approximately constant over a wide rangeof conditions with an order of magnitude of 5–10 mg cm^–2s^–1.     

THE PLACE OF A SPECIES IN THE FOREST, WITH SPECIAL REFERENCE TO WESTERN NORTH AMERICAN SPECIES OF CONIFER USED IN BRITAIN

The nature of the differences between the climates of the QueenCharlotte Islands and of England and Wales are discussed briefly.The author then examines the site tolerance of the principalspecies within their own natural range, comparing particularlythe place of Sitka spruce (Picea sitcbensis Carr.) in the foreston Graham Island with its place around Terrace and describingthe Douglas fir forests of the drier eastern side of VancouverIsland and the place that Sitka spruce has in them. Factorsaffecting the growth of Douglas fir (Pseudotsuga taxifolia Brit.)are discussed. It is suggested that care is needed before itis assumed that a markedly dry summer period such as occursin much of the Douglas fir region entails a ‘dry’period as regards supply to the tree. Illustrations are givenfrom eastern North America of changes in type of site occupiedby a species with change in climatic or edaphic environment.Observation of the relative position of a species in the forestas climate changes makes it possible to suggest the place ofSitka spruce and Douglas fir in Great Britain.     

Developments in Yield Control and Inventory in British Forestry

Forecasting timber production many years in advance has becomea problem of considerable importance in this country becauseof the need to co-ordinate the rapid increase in productionwith the establishment of new timber-using industries. Thispaper describes a change in emphasis in state forest WorkingPlan Inventory methods employed in the predominantly young,coniferous forests of Great Britain and is a development ofthe methods described in a previous paper (‘Problems ofYield Control and Inventory in British Forestry’—D.R. Johnston, 1960, Forestry, 33, 19–36). Total enumerationof the growing stock is no longer the main objective of forestinventory and the new approach is directed towards (a) assessmentof the productive potential of each subcompartment and (b) theintensive enumeration of sample or ‘index’ subcompartmentsinitially and at each re-enumeration as a check on growth predictionsand yield control.     

History of Peridermium Stem Rust of Scots Pine (Pinus sylvestris L.) in Thetford Forest, East Anglia

‘Peridermium’ stem rust (Peridermium pini (Pers.)Lev.) has been present in Thetford forest for at least 40 years,but has only become a major problem during the last decade.Survey data are presented which show that there has been a dramaticincrease in the disease in crops of Scots pine between 1964,when with less than one per cent of the trees showed symptoms,and 1979, when the figure was 10 per cent. The disease has apparentlyspread outwards from a central focus in the middle of the forest.In four plots the proportion of trees with visible symptomshas increased in five years from an average of 28 per cent to46 per cent. However, only 1–2 per cent of the trees havedied annually, and it appears that many trees with ‘deadtops’ may survive for long periods. The situation in Thetfordseems to contrast with that in north-east Scotland, where limiteddata suggest there has been no appreciable increase in diseaseincidence during recent years. There are only two records ofthe disease on Corsican pine (P. nigra var. maritima (Ait.)Melville) in Thetford.     

Determining productivity gains from herbaceous vegetation management with 'age-shift' calculations

Gains in stand volume that result from competition control andfertilization are sometimes reported as ‘percentage gains’.Because percentage gains arithmetically decline over time asstand volume increases, plantation managers have difficultyin using percentage gains to project growth and revenues. The‘age-shift’ method quantifies the year advancementsin stand growth due to silvicultural treatments and, for herbaceousvegetation management, it has been proposed that this metricis less likely to change after the juvenile growth phase. Totest the sensitivity of the ‘age-shift’ method totime and hardwood competition, we used 20-year volume data from11 loblolly pine (Pinus taeda L.) studies that had early completeherbaceous and woody competition control. Volume growth gainswere expressed in terms of percentages and ‘age-shifts’.On all sites with no woody competition, percentage gains declinedfrom age 8 years to age 20 years. In contrast, age-shift estimateson these plots either remained constant or increased over time.However, in four cases where woody basal areas were greaterthan 4 m² ha^–1 at age 15 years, age-shift gains due toherbaceous control decreased and eventually resulted in volumelosses. When evaluating the response to early herbaceous competitioncontrol, age-shift calculations have promise as a useful predictivetool on sites with low levels of hardwood competition. Fivemethods for calculating age-shift are presented.     

Effects of the extreme drought in 2003 on soil respiration in a mixed forest

We present a field study on the drought effects on total soil respiration (SR_t) and its components, i.e., “autotrophic” (SR_a: by roots/mycorrhizosphere) and “heterotrophic” respiration (SR_h: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SR_a and SR_h were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SR_a and SR_h were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T _s) and moisture sensitivity of SR_a and SR_h underneath both tree species, and (2) examined whether drought caused differential decline of SR_a between spruce and beech. Throughout the study period, SR_a of beech accounted for 45–55% of SR_t, independent of the soil water regime; in contrast, SR_a was significantly reduced during drought in spruce, and amounted then to only 25% of SR_t. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l⁻¹ a⁻¹), but remained at levels similar to those in 2002 in beech (about 470 mg l⁻¹ a⁻¹). This species-specific root response to drought was related to a stronger decline of SR_a in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SR_a and SR_h to changing T _s and available soil water was stronger in SR_a than SR_h in spruce, but not so in beech. It is concluded that SR_a determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands."     

The lichens of the Scottish native pinewoods

During the last 30 years, 434 epiphytic lichen taxa have beenrecorded from the native pinewoods of Scotland. Of these lichens,about 219 have been recorded from the bark, lignum or stumpsof pine (Pinus sylvestris) itself. Eighteen species are, inthe British Isles, confined to these native pinewoods, whichare themselves confined to the Scottish Highlands. A resumeis presented of the habitats and niches available to lichensin the native pinewoods, using examples of the most characteristicspecies for each. The ‘pinewood habitat’ for epiphyticlichens is by no means restricted to pine, but also includesassociated trees, shrubs and subshrubs, especially birch, rowan,holly, aspen, juniper and heather. The strong east–westvariation in the composition of the lichen flora is outlined.By comparison of species lists from pinewood stands of knownancientness with those from mature plantations distant fromthe ‘core’ pinewoods, an additional c. 60 speciescan be considered as indicative of an ecological continuityof the pinewood habitat. From this pool of c. 77 species, aNative Pinewood Index of Ecological Continuity is presentedas a useful tool for biodiversity assessments of pinewoods.     

Development of the Structural Root System of Sitka Spruce

Growth ring analysis was carried out on root systems of Sitkaspruce trees which had been planted 8 and 34 years previously.Retrospective measurements were made on root extension, andon cross-sectional area increment near the root base. The extensionof main lateral roots started slowly, then increased, and theyachieved a mean length of 4.4 m in 8 years. Differentiationinto roots of widely different radial growth rates took placeduring the first 6 years, resulting in 3 to 11 ‘major’woody roots and a large number of small ‘minor’ones, with some of intermediate vigour, radiating from the stump.The major roots established during the first few years constitutedthe main structural root system at 34 years. Many of the minorroots stopped growing in diameter after a few years, but werestill alive and extending at 34 years. The differentiation intomajor and minor roots is discussed with reference to their originon the root system, primary xylem diameter and the local environment.     

The Detection of Viruses with Nematode Vectors in Six Woody Hosts

Arabis mosaic virus (AMV) was transmitted to herbaceous hostswith foliar sap from the following donor woody plants: Fraxinusamericana L. with distorted leaves having chlorotic ring patterns,Hedera helix L. with foliar yellow chevrons, rings or blotches,Jasminum officinale L. ‘aureo-marmorata’ Hort. withyellow rings and blotches, Laburnum vossii Hendr. with stuntingshoot proliferation and terminal dieback (button-top) and Spiraeadouglasii Hooker with leaf enations and yellow veins. In threeinstances, affected donors plants were growing in soil infestedwith nematodes carrying virus. Strawberry latent ringspot virus with AMV was transmitted fromone of the H. helix donors; AMV and tobacco rattle virus weretransmitted from roots but not leaves of Populus x euramericana.     

Sitka Spruce and Douglas fir Seedlings in the Nursery and in Cold Storage: Root Growth Potential, Carbohydrate Content, Dormancy, Frost Hardiness and Mitotic Index

Seedlings (transplants) of 2+1 Sitka spruce (Picea sitchensis(Bong.) Carr.) and 1 + 1 Douglas fir (Pseudotsuga menziesii(Mirb.) Franco) were grown in a nursery at the Bush Estate,Scotland. Batches were lifted and cold stored at 0.5°C inNovember, December and January. Changes in growth, shoot apicalmitotic index, root growth potential (RGP), carbohydrate content,bud dormancy and shoot frost hardiness were monitored throughoutthe winter by taking samples at intervals from the nursery andfrom cold storage. Frost hardening occurred during the later stages of bud development(as mitotic indices decreased); autumn hardening was arrestedwhen seedlings were put in cold store, and some dehardeningoccurred in cold storage, especially in spring. Bud dormancystarted, and was greatest, just after bud growth (mitotic activity)virtually ceased; chilling in cold store was almost as effectivein releasing dormancy as natural chilling. The concentrationof total nonstructural carbohydrates stayed more or less constantat 100–150mg g^–1 from September to April in thenursery; in cold storage carbohydrates were depleted at 0.4–0.6mgg^–1 d^–1 (corresponding to respiration at 0.03–0.05mgCO₂ g^–1 h^–1) until there was only 40–50mgg^–1. Root growth potentials in the nursery increased in December,once the buds ceased growth, became dormant and had receivedsome chilling. Sitka spruce was ‘storable’ in November,before RGPs increased, but they then failed to achieve maximalfrost hardiness or ROP. Winter RGPs were high in Sitka spruceand were increased or maintained in cold storage, whereas RGPswere low in Douglas fir and decreased immediately after storage(except when stored in January). By the end of April, the RGPof cold stored Sitka spruce was much higher than that of directlifted plants. ROP changes in the nursery and in cold storagewere not consistently related to changes in seedling carbohydratecontents, shoot frost hardiness or bud dormancy. In practical terms, it was concluded that (1) the optimum dateto start lifting bare- rooted conifer transplants in the autumnis when their shoot apical mitotic indices have decreased tonear zero, and their RGPs have risen sharply; (2) high RGPsmay depend as much on the morphology of the roots (e.g. numberof undamaged root apices) as on the physiology of the shoots(e.g. carbohydrate status, dormancy and frost hardiness); and(3) in spring, transplants kept in cold storage since November,December or January are more frost hardy, slightly more dormant,and (in May) have higher RGPs than transplants lifted from thenursery.     

ERRATUM

THE author has called our attention to a mistake in her paper‘Squirrel Populations and their Control’ publishedin the recent Supplement to Forestry ‘Wildlife in theForest’. The mistake occurs in the second paragraph ofpage 19 where the fall in the estimated population of grey squirrelsfollowing a mast failure should be 468 to 66 and not 468 to466.     

Floristic Diversity in Some Central Swedish Forests

The species of vascular plants present in 0·5 ha sampleplots were recorded in 19 areas of forest planted on formeragricultural land, 21 areas of old or ‘virgin’ forestand 102 areas of ‘ordinary forest’ in east centralSweden. In the case of the planted forest areas, the main correlationwith floristic diversity was with the amount of daylight reachingthe forest floor. In the ‘old forest’ and ‘ordinaryforest’ sites, however, the main correlations were withthe pH value of the surface soil, the amount which this variedwithin the sample plot, and the proportion of broadleaved treesin the tree canopy; these factors were inter-correlated witheach other to some extent. The ‘old forest’ sites,which had the most acidic surface soil and least diverse flora,did not appear to be serving any particularly useful functionin terms of conservation of the flora, at the present time.It is concluded that, if a diverse forest flora is requiredfor any particular reason, the complete closure of the canopyshould be avoided. A mixture containing some broadleaved treesis likely to be preferable to a pure crop of pine or spruce.There appears to be, however, considerable scope for the developmentof a diverse flora within timber-producing forests consistingmainly of spruce and pine.     

Shading Reduces both Visible and Invisible Frost Damage to Norway Spruce Seedlings in the Field

Two-year-old cuttings of Norway spruce were subjected to nightfrosts in spring on an exposed site in southern Sweden. Shadingwas used to assess the influence of sunlight on the extent ofdamage resulting from night frost. Chlorophyll fluorescencewas measured in needles in flushing shoots, and in shoots atthe stage of bud burst. The F_v:F_m ratio was significantly lowerfor plants exposed to light, compared with shaded plants onthe days following the night frost (minimum temperature –6°C).The effect was similar both in 1-year-old and current year needles.The low F_v:F_m ratios indicate damage to photosystem H, causedby an interaction between sub freezing temperatures and highlight intensity. Shading also increased the survival of flushingshoots. It is suggested that regeneration of Norway spruce onsites exposed to frost should be carried out in partial shade,for example under a shelterwood.