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Changes in Whole-Tree Water Use Following Live-Crown Pruning in Young Plantation-Grown Eucalyptus Pilularis and Eucalyptus Cloeziana

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Changes in Whole-Tree Water Use Following Live-Crown Pruning in Young Plantation-Grown Eucalyptus Pilularis and Eucalyptus Cloeziana Forests 2013, 4, 106-121; doi:10.3390/f4010106 OPEN ACCESS forests ISSN 1999-4907 www.mdpi.com/journal/forests Article Changes in Whole-Tree Water Use Following Live-Crown Pruning in Young Plantation-Grown Eucalyptus pilularis and Eucalyptus cloeziana Philip J. Alcorn 1,2, David I. Forrester 3,4,*, Dane S. Thomas 5,6, Ryde James 1, R. Geoff B. Smith 5,7, Adrienne B. Nicotra 2 and Jürgen Bauhus 3 1 Fenner School of Environment and Society, The Australian National University, Canberra, ACT 0200, Australia; E-Mails: [email protected] (P.J.A.); [email protected] (R.J.) 2 Research School of Biology, The Australian National University, Canberra, ACT 0200, Australia; E-Mail: [email protected] 3 Chair of Silviculture, Faculty of Environment and Natural Resources, Freiburg University, D-79085 Freiburg, Germany; E-Mail: [email protected] 4 Cooperative Research Centre for Forestry, Private Bag 12, Hobart 7001, Australia 5 Forests NSW, PO Box J19, Coffs Harbour NSW 2450, Australia; E-Mails: [email protected] (D.S.T.); [email protected] (R.G.B.S.) 6 South Australian Research and Development Institute (SARDI), Climate Applications, GPO Box 397, Adelaide SA, 5001, Australia 7 Forest Science Centre, School of Environment, Science and Engineering, SCU Lismore NSW 2480, Australia * Author to whom correspondence should be addressed; E-Mail: [email protected]; Tel.: +61-761-203-8628; Fax: +61-761-203-3781. Received: 28 November 2012; in revised form: 23 January 2013 / Accepted: 1 February 2013 / Published: 5 February 2013 Abstract: Pruning of live branches is a management option to enhance wood quality in plantation trees. It may also alter whole-tree water use, but little is known about the extent and duration of changes in transpiration. In this study, sap flow sensors were used to measure transpiration for 14 days prior to, and 75 days following the removal, through pruning, of the lower 50% of the live-crown length of 10–11 m tall four-year old Eucalyptus pilularis Sm. and E. cloeziana F. Muell. trees. Pruning had no effect on stem growth, sapwood water content or radial pattern of sap velocity in either species. Pruning Forests 2013, 4 107 reduced mean daily water use by 39% in E. pilularis and 59% in E. cloeziana during the first eight days after pruning. Thirty six days after pruning there were no longer any significant differences in transpiration rates between pruned and unpruned trees in either species. Our results show that pruning of live branches had only a short-term effect on whole-tree transpiration in these sub-tropical eucalypt species. Keywords: defoliation; eucalypt; compensation heat pulse technique; sap flow; sapwood 1. Introduction Whole-plant water use is influenced by soil nutrient and moisture conditions [1–4], humidity of air adjacent to leaves [5,6] and the supply of water from conducting stem tissue [7]. Rapid reductions in functional or effective photosynthetic leaf area following shading, disease or defoliation have the potential to dramatically alter whole-plant water use, although not necessarily in proportion to the leaf area reduction [8,9]. For example, tree conductance was immediately reduced following shading of the lower 78% of foliage of a Pinus radiata tree [9]. However, when tree conductance was normalised with respect to the illuminated foliage area, water use actually increased by 50%–75%. Similarly, the removal of about 45% [10] or 60% [11] of the leaf area of Eucalyptus globulus tree crowns resulted in higher transpiration per unit leaf area. Pruning 75% of the leaf area of Eucalyptus nitens trees reduced transpiration of the dominant 200 trees ha−1 by about 16%, but increased transpiration per unit leaf area, 2–3 years after pruning [12]. Enhanced water use by the remaining foliage following a reduction in total photosynthetic foliage area can be partially attributed to short-term compensatory changes in leaf-level processes. Pruning increased rates of stomatal conductance to water vapour or instantaneous rates of transpiration of retained leaves after pruning in Eucalyptus globulus or Eucalyptus nitens [12–14]. Two shading studies demonstrated large increases in stomatal conductance in the illuminated foliage following a reduction in photosynthetic leaf area by shading the lower foliage, and these effects were fully reversed when shade was removed [8,9]. The magnitude and duration of whole-plant water use reduction following a reduction in total photosynthetic leaf area is influenced by plant architecture and the severity of the change in leaf area. Whole plant and leaf-level compensatory responses differ depending on the position of functional leaf area removal in the plant crown [2,8,9], the proportion of the leaf area removed [13,15] and the frequency of removal [16]. The duration of whole-plant water use reductions following reductions in photosynthetic leaf area depends on the rate of leaf area production in the crown [17]. In fast-growing species with rapid turn-over of leaf area [18], including eucalypts, reductions in water use following defoliation may be relatively short-lived under favourable environmental and climatic conditions. However, such reductions may be useful during short-term periods of low rainfall to reduce the mortality rates and the susceptibility of trees and stands to water stress. Removal of live branches (live-crown pruning) is a form of artificial defoliation. It is employed to enhance timber quality in tree plantations. By removing the lower branches from the stem, knots and branch-related defects can be restricted to a central knotty core of the stem and high-value knot-free Forests 2013, 4 108 timber can subsequently be produced in the pruned zones. The compensation heat-pulse method (CHPM) was used to measure transpiration before and after moderate live-crown pruning in two commercially important timber species, Eucalyptus cloeziana F. Muell and E. pilularis Sm. Both species are grown in plantations in the sub-tropical region of southern Queensland and northern New South Wales, Australia, but contrast in shade tolerance and crown architectures, such that E. pilularis is less shade tolerant and has smaller crowns [19]. We hypothesised that live-crown pruning would lead to an initial reduction in whole-tree transpiration, but that this response would be short-lived. While live-crown pruning is of interest from a timber management perspective, the findings of this study are also directly applicable to natural plant defoliation (herbivory). 2. Materials and Methods 2.1. Study Site and Treatment Design The pruning trial was established in a research plantation near Nana Glen in northeastern New South Wales (30°1′S, 153°8′E), where both species were planted in adjacent stands. This well-drained site contains gently sloping deep (1–1.5 m) brown and yellow earths soils [20] derived from late carboniferous siltstone, mudstone and conglomerate [21]. The site is approximately 165 m above sea level and receives a moderately high annual precipitation of 1437 mm (measured 1920–2004), distributed with a distinct winter minimum and summer/autumn maximum. Original site vegetation consisted of a tall open mixed hardwood forest including E. pilularis, E. intermedia R. Baker and E. microcorys F. Muell. The site was cleared early last century and subsequently converted to pasture for grazing. To prepare the site for planting, the soil was ripped in a north-south direction in September 2000 to a depth of 0.7 m and mounded to obtain 4 m wide row spacings. A second cultivation was completed one month later to decrease soil tilth for planting. Herbicides (glyphosate 4 L ha−1, simazine 2.5 kg ha−1and metolachlor 1.5 L ha−1) were applied to mounded soil one month prior to planting in December 2000. Eucalyptus pilularis (Whian Whian State Forest seedlot) and E. cloeziana (Pomona State Park and Mebbin State Forest plantation seedlot) were planted at a stocking of 1250 trees ha−1. Each seedling was fertilised with 9 g elemental nitrogen and 10 g elemental phosphorus in the form of diammonium phosphate at the time of planting. Post-plant weed control involved applications of haloxyfop (0.5 L ha−1) and clopyralid (0.8 L ha−1) to planted mounds one and four months after planting. Six 30 × 30 m blocks of each species were located on a gentle east-facing slope (<4°). Within each block, two trees were selected with a similar diameter at breast height (1.3 m, DBH). These 12 trees per species were of the co-dominant or dominant crown class within the stand [22], straight and single stemmed, free of visible health defects and surrounded by four immediate neighbours in each direction. On the 6 October 2004, one randomly selected tree in each block was pruned to remove the lower 50% of the length of the live-crown. Live-crown length was visually defined as the distance between the tree height and the stem insertion height of the lowest live branch contained within a geometrically regular crown envelope [23]. In practice, pruning regimes for these species are being developed but it is likely that all pruning up to a height of about 5–6 m would be completed by about age five or six years. Forests 2013, 4 109 2.2. Sap Flow Measurements The twelve selected trees per species were monitored for 87 days with sixteen sap flow sensor units (Model SF100 Greenspan Technology, Warwick, Queensland, Australia). Units were deployed for 12 days before pruning (24 September to 5 October 2004) and 75 days after pruning (7 October to 20 December 2004) using a roaming sensor technique [24–26]. Trees were not monitored on the day (6 October 2004) on which pruning occurred.
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