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Original article An evaluation of decapitation as a method for selecting clonal Quercus petraea (Matt) Liebl with different branching intensities R Harmer, C Baker Forestry Commission Research Station, Alice Holt Lodge, Wrecclesham, Farnham, Surrey, GU10 4LH, UK 2 25 (Received September 1994) January 1993; accepted Summary — The effect of decapitation on branch production in 5 clones of oak was observed over the 2 flushes of growth occurring during 1 season. Concurrent experiments were carried out under natu- ral conditions in the nursery and 2 different temperature regimes in growth chambers. Decapitation had no effect on the number of buds becoming active but usually increased both the proportion of active buds forming branches and the number of branches produced during each flush. More branches were formed during the first flush of growth but the largest effect of decapitation occurred during the second flush. There were significant differences between clones but the clonal order of branchiness varied between flushes and treatment. Lower temperatures reduced the rate of shoot development but had only small effects on the length of new leading shoot and the proportion of buds becoming branches. The significance of these results for the selection of oaks with different branching patterns is dis- cussed. Quercus petraea / clone / bud / branching Résumé — Une évaluation de la décapitation comme méthode de sélection clonale de Quer- cus petraea (Matt) Liebl présentant différentes intensités de branchaison. Les effets produits par la décapitation sur la ramification observée sur 5 clones de chêne ont été étudiés au cours des 2 vagues de croissance se produisant en une saison. Des expériences sont effectuées simultanément en pépinière, dans des conditions naturelles, et en laboratoire, en ayant recours à 2 régimes de tempéra- tures différents. La décapitation n’affecte en rien le nombre des bourgeons devenant actifs (tableau III), alors qu’elle augmente généralement à la fois la proportion de bourgeons actifs formant des branches et le nombre de branches produites pendant chaque vague de croissance (fig 4). Bien que la ramifica- tion soit plus fréquente sur le dernier cycle de l’année précédente que sur le premier cycle de l’année en cours, la décapitation a une plus grande influence sur la ramification dans le second cas que dans le premier (figs 5 et 6). Des variations significatives apparaissent d’un clone à l’autre, mais aussi selon le cycle de croissance considéré et selon les traitements (tableau IV). Il s’avère que les températures plus basses réduisent la vitesse de développement de la pousse, mais n’ont que peu d’effet sur la longueur
de la nouvelle pousse apicale et la proportion de bourgeons donnant naissance à des branches (figs 2 et 3). La portée qu’ont ces résultats sur la sélection des chênes présentant des systèmes de ramifica- tion différents fait l’objet de discussions. chêne / clone / bourgeon/ ramification Quercus petraea / or large numbers of INTRODUCTION trunk; large branches, branches will significantly reduce the qual- ity and hence value of oak timber. Careful Deciduous oaks are some of the most silvicultural practice can be used to manip- important hardwood timber trees in north- ulate branching but the normal tendency of temperate Europe and, for example, in Great oak to produce a spreading crown with large Britain they form 30% of broadleaved high branches is difficult to suppress whilst main- forest providing 25-30% of hardwood timber taining an acceptable combination of height for sawmills (Evans, 1984). However, the and diameter growth. An important part of quality of oak timber is very variable and our oak improvement programme aims to there may be a 10-fold difference in the gain a better understanding of the develop- value of high- and low-grade timber (White- ment and control of branching and identify man et al, 1991). Despite the commercial genotypes with superior stem and crown importance of oak there has been little form. emphasis on improvement of planting stock Studies with obeche (Triplochifon scle- by the selection of superior genotypes and K Schum), a fast growing tropical large scale trans-European provenance trials roxylon, tree, have shown that it is possible to relate with Quercus robur L and Q petraea (Matt) Liebl are only just beginning. These will not branching in small, young, clonal plants to yield final results for several decades and that of larger plants growing in the field. the uncertainty of seed supply may, even When small plants were decapitated, the then, prevent use of the best provenances. number of branches produced varied Several studies have shown that it is pos- between clones (Leakey and Longman, sible to produce clonal oaks either by micro- 1986); clonal field trials showed that after 5 propagation of softwood cuttings (Klein- years’ growth the number of branches on schmit et al, 1975a; Spethmann, 1986; the main stem was positively correlated Meier-Dinkel, 1987; San-Jose et al, 1990). with branch production in decapitation Such procedures could be used to supply experiments (Leakey and Ladipo, 1987). suitable planting stock and avoid the The following experiments were carried out vagaries of seed supply. At present these in order to evaluate the use of decapitation methods are only successful with some juve- as a method for selecting oaks with differ- nile material but there is no current method ent branching patterns. Growth in oak is for determining whether the juvenile clones determinate and there are 1 or more dis- capable of mass propagation will produce crete periods of shoot extension during the high quality trees. The UK Forestry Com- growing season which are, in part, under mission’s oak improvement programme is endogenous control (Barnola et al, 1986; investigating methods of identifying supe- Alatou et al, 1989; Barnola et al, 1990; Par- rior trees when they are juvenile and can mentier et al, 1991; Barnola et al, 1993). be used for clonal propagation. As the formation of lateral branches The quality of oaks for saw logs is related appears to differ between periods of growth to the size and number of branches on the occurring at different times of the year
required and given liquid fertiliser (N:P O, :K 5 O 2 (Harmer, 1992b), experiments were car- 8:4:4) at 14-d intervals. During the first 6 weeks of ried out using overwintered shoots and the experiment, leaves on some plants in the those produced during the first period of warm environment developed mildew; these growth in spring. leaves were removed immediately. No mildew developed on plants in the cool chamber. The few aphids that appeared were controlled by hand METHODS during experimental observations. Plants in the nursery were sprayed with pyrethrum-based insec- ticide and sulphur to control aphids and mildew, respectively. Plant culture and experimental treatments Assessment 1989 leafy cuttings were taken During summer from shoots growing on stumps of 10-year-old Q petraea trees felled during winter 1988. Cuttings The plants in the growth chambers were observed were rooted using methods described by Harmer on alternate days throughout the experiment, and Baker (1991). Surviving cuttings were over- which lasted for 2 periods of shoot growth. Three wintered in the trays of substrate used for rooting sections of leading shoot were observed during and then grown outdoors for 1 season in 10 cm the experiment (fig 1): a) original shoot the — plastic pots containing 3:1 peat/grit compost with terminal section of shoot produced by the sec- slow release fertiliser (18:11:10, N:P 4.3 O, :K 5 O 2 ond period of growth in 1990, this carried over- kg m ). -3 wintering buds; b) first-flush shoot the section — produced during the first period of growth in the In February 1991, similar sized plants from experiment; and c) second-flush shoot the 5 clones from parents with apparently different — section produced during the second period of growth form were repotted into 12.5 cm diameter growth in the experiment. For decapitated plants plastic pots of compost. The plants selected had the leading shoot was defined as the longest produced 2 flushes of growth in 1990 and had branch which grew from the lateral buds at the tip a live terminal bud. Most plants had produced of the shoot. 1-2 branches which were removed after repot- ting. Plants were then randomly assigned to 2 The times taken to reach the following states decapitation treatments in 3 environmental con- of development were scored for the most ditions; there were 5-10 plants of each clone advanced bud on the original shoot during the receiving the decapitation treatments in each first period of growth: d) bud expansion green — environment. areas appearing between bud scales but no i. the terminal bud removed, leaves visible, buds which reached this state were Decapitation was — using forceps, from half of the plants at the start of regarded as active; e) first visible leaf beginning — both the first and second flushes of growth; the of bud opening and shoot extension; f) leaf expan- remaining plants were untreated, intact, controls. sion new shoot no longer extending, leaves — expanding; and g) end of flush leaves fully ii. Environment equivalent numbers of each — — expanded. The same features, except (e), were clone receiving the 2 decapitation treatments assessed for buds on the first-flush shoot during were grown in growth chambers under 2 different the second period of growth. During both periods temperature regimes: warm, 20°/15° day/night; of growth the total number of buds active was cool, 15°/10° day/night. Plants were also grown assessed at 8-d intervals. under natural conditions in the nursery. At the end of the first period of growth the num- Environmental differences between chambers ber of lateral branches on the original shoot was were minimised: day length was 18 h and sup- counted before removing all new growth except plied by both fluorescent tubes (Sylvania, Cool the leading shoot. During the second period of white) and tungsten lamps; photosynthetically growth a few buds became active on the original active radiation at canopy height was adjusted weekly to 145 μmol m s day/night water -2 - shoot, these were not counted. After completion 1; of the second period of growth the number of lat- vapour pressure deficits were approximately 2.3/1.0 mb, respectively. Pots were watered as eral branches on the first-flush shoot was counted
and the lengths of the leading shoots produced RESULTS during both the first and second periods of growth measured. significant effects of clone and There were Plants growing under natural conditions in the decapitation on the branching of plants but, nursery were treated in the same way as those in with the exception of rate development, the the growth chambers but only shoot lengths and branch numbers were assessed. effects of temperature were small (table I, fig 2). The presence or absence of the terminal After the mother trees had been felled, mea- surements were made of the length and number bud had no significant influence on the time of branches on the final 3 sections of shoot pre- taken to reach each stage of development sent on the leader and the 4 major crown therefore figure 2 shows the means of data branches. These sections, which were equiva- over both decapitation treatments. There lent to those of the experimental plants, are also were significant differences between clones termed original, first-flush and second-flush shoots and between temperature conditions in the (fig 1). number of days taken for the most advanced bud to reach each stage of development. Overwintered buds on clones in the warm Statistical analysis and presentation chamber reached bud expansion in about of data 11 d and finished their development after 26 d, the second period of growth started Due to the large differences in experiment times at day 54 and finished 10 d later. Plants in and conditions, data for plants grown in the growth the cool growth chamber developed more chambers, the nursery and the field have been slowly; the first period of growth lasted for 42 analysed separately. The effects of clones and treatments were investigated by analysis of vari- d and the second period started at day 79 ance. As previous studies have shown that bud and lasted 25 d. The rate of development and branch numbers are related to shoot length of plants growing under natural conditions (Harmer, 1989a, 1992a) analyses of these data was slower than that for either chamber. used length as a covariate; any levels of signifi- Expansion of overwintered buds began in cance given in the text, tables or figures result from these analyses. However, the means and standard errors of differences between means presented in tables and figures are not adjusted for the covariate.
about 3.5-5-fold longer than the first-flush the last week of March, the first period of growth being completed by the end of May shoots and there were significant differ- after about 70 d; the second period of growth ences between clones (p ≤ 0.001), the started in June and ended in July. This mean length varying between 50 and 175 observation is similar to those describing mm for clones 7 and 4 respectively. For the normal pattern of growth under natural both the first-flush and the second-flush conditions. leading shoots the rank of clones according to length was clone 7 < 10 < 5 < 2 < 4. For plants in the growth chambers, decapitation had no significant effect on Length data for the plants grown under length of the leading shoot produced and natural conditions are presented in figure whilst lower temperatures reduced the 3b. Overall trends between flushes were length of the second-flush leading shoot by similar to those for plants grown in cham- between 6 and 30%, the effect was only bers: the first-flush shoots were the shortest significant at the 5% level (table I). The and second-flush usually the longest, but mean lengths of the leading shoots pro- shoots were generally shorter and the rank duced by each clone during each period of order of clones differed. growth over all treatments are shown in fig- The mean lengths of the shoots present ure 3a. The mean length of the original on the mother trees were always greater than shoot varied between 37 and 50 mm and those on the clonal plants (table II; fig 3a,b). did not differ significantly between clones. Whilst the first-flush shoots of these trees For all clones the mean length of the first- were usually shorter than the original shoots flush was always smaller than the original the difference between second-flush and orig- shoot; clone 7 was the shortest and clone 4 inal was less obvious than for the clonal the largest, at 14 and 37 mm respectively (fig 3a). The second-flush shoots were plants.
for control and decapitated plants, respec- tively (fig 4). Plants in the cool chamber produced approximately 25% more branches on original shoots than those in the warm chamber (p≤ 0.05). Analysis of the data for the first-flush shoots showed significant interactions between clones, ter- minal and temperature treatments (table I) which were due to clones 5 and 7 that showed a less obvious or opposite response to decapitation at the different temperatures. The numbers of branches present on each shoot are shown in figures 5 and 6; as the only effect of temperature was a small 3-way interaction (table I), the data for both warm and cool chambers have been com- bined (fig 5). There were no sylleptic branches. For plants under all conditions there were significant differences between clones in the number of branches formed on each shoot. In general, the original shoots carried more branches than first- flush shoots and over all clones and treat- ments the mean number of branches pre- sent on a shoot varied from 0 to 5.75; these values were found for clone 4 grown under natural conditions (figs 5 and 6). The effects The numbers of buds that became active of decapitation were usually positive with growth chamber plants during the 1st on the largest percentage increases in num- and 2nd period of growth are shown in table bers of branches occurring after decapitation III; these were not influenced by decapitation of the first-flush shoots (figs 5 and 6). Decap- or temperature (table I). More buds became itation caused increases of 0-140% in the active on the original shoot than on the first- number of branches on original shoots and flush shoot, the number varied between 10-560% on first-flush shoots. The only 6.8-9.8 and 4.9-5.8, respectively. exception was clone 5 growing under natu- ral conditions, where decapitation caused Both clone and decapitation had signif- a 60% reduction in number of branches on icant effects on the proportion of active buds the first-flush. On mother trees the original that became branches on original and first- shoots also carried the most branches and flush shoots (p≤ 0.001) (table I; fig 4). For in general each shoot had more branches intact, control plants the proportion of active than comparable control, clonal plants (table buds forming branches on the original shoot II, figs 5 and 6). varied from 0.13 to 0.42 (fig 4), decapitation increased this to between 0.33 and 0.60. In order to compare the branchiness of The proportions of active first-flush buds each shoot it was necessary to allow for the large differences in length by calculating forming branches were similar to these, number of branches per unit length of shoot ranging between 0.08-0.55 and 0.29-0.60
In most cases, control trees were (table IV). DISCUSSION less branched than corresponding decapi- tated plants under the same growing con- These investigations showed that decapi- ditions, and shoots on mother trees were tation stimulated lateral branch production nearly always less branched than experi- but the magnitude of the response varied mental plants; the difference between between clones. Although the influence of flushes was less marked. The number of decapitation was the same for each section branches per millimetre varied between of shoot there appeared to be quantitative zero, for control first-flush shoots of clone differences between original and first-flush 4 under natural conditions, and 0.335 for shoots that varied with growth conditions. the original shoots of clone 2 receiving the Differences in response between these same treatment. The rank numbers of the shoots was probably related to their physio- clones according to branchiness for each logical state reflecting the differences treatment are also presented in table IV. between acrotony (apical control) and apical Although the original shoot of clone 7 was dominance (Brown et al, 1967; Champagnat generally the least branched, the rank order et al, 1971; Champagnat, 1978; Crabbé, of the clones depended on treatment. There 1987; Champagnat, 1989). Original shoots was no obvious relationship between branchiness of the experimental plants and were leafless, with new shoot growth devel- the mother trees. oping from 6-month-old buds emerging ca
sistent with those of Leakey and Longman winter dormancy. In con- from period of a (1986), who found that temperature had lit- trast, first-flush shoots leafy, actively were growing and their new shoots developed tle effect on percentage bud activity, the from buds that had experienced only a short influence of temperature on branching is period of rest. unclear. Most studies of apical dominance have been with herbaceous plants and Casual observations of seedlings grow- results on the influence of temperature on ing in the nursery and greenhouse, and both these and woody plants are inconclu- shoots developing within and outside sive. There are a number of studies which treeshelters (Potter, 1991) had suggested show that lower temperatures can reduce that temperature was an important factor apical dominance and increase branching influencing branching. However, results (Bollman et al, 1986; Rosa, 1986; Moe, from plants growing in the controlled envi- 1988) but there are others which show the ronment chambers suggest that the effects opposite or no effect (White and Mansfield, of temperature are relatively small com- 1978; Struik et al, 1989). In the experiments pared to other factors. Low temperature described using oak, only 1 chamber was had the predictable effect of reducing rate used for each temperature and any effects of development (fig 2) but had few other ascribed to temperature may be due to significant effects which were most often other unknown differences in conditions apparent as interactions with other factors between chambers. Further experiments (table I). Although these results were con-
needed to define more precisely the of growth. Comparison of this result with are role of temperature in branching and growth those for other temperate trees is difficult to of oak. do, not only because the results of prun- ing experiments are very variable, depend- The relative lengths of shoot produced ing on many factors including vigour, grow- during the first and second periods of growth ing conditions, time of treatment and plant by each clone was typical of oak. The first- age (Mika, 1986; Crabbé, 1987), but also flush shoot is usually shorter than the sec- because the pattern of growth shown by ond-flush shoot produced by recurrent flush- oak is different from that for temperate fruit ing during summer in both field and nursery grown plants (Dostal, 1927; Gruber, 1987; trees for which most information is avail- Harmer, 1992b). The reasons for this are able. In general, dormant pruning of tem- unknown but may be due to a better sup- perate fruit trees stimulates the develop- ply of mineral nutrients and carbohydrate ment of longer shoots (Mika, 1986) but as to the buds from plants with active roots and these grow more or less continuously when leaves compared to the leafless original conditions are favourable they are not com- shoot. parable to oak shoots which grown rhyth- mically even when conditions are ideal. Although these experiments found that The length of new leader produced by lat- length of shoot varied with clone, decapi- tation had no effect on length of the new eral buds on decapitated plants was not leading shoot produced during each period significantly different from that for terminal
ship with numbers of buds and branches buds on untreated plants. This was unex- (Ward, 1964; Harmer, 1989a; 1992a). Sim- pected as the terminal bud is usually the ilar adjustments were needed in a study of largest on a shoot and it suggests that branching in poplar clones (Sauer, 1959). there may be little relationship between bud size and shoot length and that bud Decapitation had no effect on the number and shoot growth is strongly influenced by of buds that became active but significantly correlative effects. increased both the proportion of buds that became branches and the number of Studies of branch production following branches on the shoot (table I). The great- have also been made on tem- decapitation est percentage increases in branch num- perate fruit trees where pruning usually ber occurred for first-flush shoots. These increases the number of lateral branches results reflect the pattern of apical domi- produced (Barlow and Hancock, 1962; Mika nance and control shown by oak (Brown et 1986). Similar results have been found for al, 1967; Champagnat, 1989): the buds on Morus alba L (Suzuki et al, 1988), Q rubra L the overwintered original shoot are under (Ward, 1964) and T scleroxylon (Leakey weak apical control and many are able to and Longman, 1986). Before investigating form branches without decapitation; in con- the differences in bud activity and branch trast, buds on the first-flush are suppressed production between clones of oak it was by strong apical dominance which is necessary to allow for shoot length which removed by decapitation. As most new was very variable and has a close relation-
branch production in the field occurs dur- II): the data in table IV show that there was ing spring any selection test should proba- obvious relationship between young no bly be based on the observation of branch- clonal plants and mother trees. ing on the original shoot; unless the The experiments described, which inves- behaviour of original and first-flush shoots tigated the relationship between genotype, can be correlated this will restrict study to branching and temperature, were made in annual observations. order to evaluate the use of decapitation There have been few studies on growth as a method for selecting clonal oak with using clonal oak and this is the first report of different branching characteristics. Only a an experiment that has investigated branch- small number of clones were used but ing in young clonal material generated from results show that the rank order of clones cuttings. Recent studies of clonal Q rubra according to branchiness varies with grow- derived from split embryos have shown sig- ing conditions and period of growth nificant differences in height and stem diam- observed. Reasons for this difference are eter between clones from a number of fam- not known but they probably result from ilies (Kolb and Steiner, 1989). Similarly, the variation in endogenous features such as length of the new leading shoots also var- those investigated in detailed physiologi- ied considerably amongst the small num- cal studies (Alatou et al, 1989; Champag- ber of Q petraea clones investigated in the nat, 1989; Parmentier et al, 1991).Simi- study described, which suggests that it larly it is not known if the branching could be possible to select clonal oaks that differences found will remain throughout show differences in rate of height growth. the life of the tree or precisely how they will Studies of 15-20-year-old grafted plants of change as the plant grows giving rise to Q robur and Q petraea have shown that mature plants with patterns of branch there can be significant clonal differences in growth and crown architecture which dif- form, crown shape, branchiness, stem fers from that of juveniles (Bartelemy et al, branch angle and tree from (Kleinschmit et 1989; Edelin, 1991).Although it may be al, 1975b) but it is not known how the form possible to develop a ’predictive test’ for of these plants developed. In contrast, the branching that is based on decapitation, it experiments with clonal Q petraea is clear that the experimental environment, described in the present study have shown physiological stage and age of the plant significant clonal differences in shoot length need to be closely defined (Harmer, 1989b). and branch production, but the final form In addition, in order to relate experimental of these clones in the field is not yet known. results to growth in the field, it will be nec- In order to understand the relationships essary to establish field trials where obser- between the growth of young clonal plants vation of branching takes place over several and mature trees in the field, it will be nec- years. However, screening even a few essary to establish trials such as that clones is expensive and even if a suitable described by Leakey and Ladipo (1987), predictive test can be developed it may be who found that the number of branches pro- impossible to screen large numbers of duced by clones of T scleroxylon in the field clones. We have established stock hedges was strongly correlated with bud activity and intend to produce sufficient plants from after decapitation of small plants in a ’pre- a variety of clones to establish field trials dictive test’. At present, the only data avail- and continue exerimental observations of able for the comparison of young and field branching in order to gain a better under- grown clones are for the final 3 sections of standing of plant growth and develop a shoot on branches of the mother trees (table method of selecting superior clones.
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