P e r t a n i ka J o u r n al of TROPICAL AGRICULTURAL SCIENCE Vol. 32 (2) Aug. 2009 A scientific journal published by Universiti Putra Malaysia Press Journal of Tropical Agricultural Science About the Journal Pertanika is an international peer-reviewed journal devoted to the publication of original papers, and it serves as a forum for practical approaches to improving quality in issues pertaining to tropical agriculture and its related fields. Pertanika Journal of Tropical Agricultural Science which began publication in 1978 is a leading agricultural journal in Malaysia. After 29 years as a multidisciplinary journal, the revamped Pertanika Journal of Tropical Agricultural Science (JTAS) is now focusing on tropical agricultural research. Other Pertanika series include Pertanika Journal of Science and Technology (JST) and Pertanika Journal of Social Sciences and Humanities (JSSH). JTAS is published in English and it is open to authors around the world regardless of the nationality. It is currently published two times a year, i.e. in February and August. Goal of Pertanika Our goal is to bring the highest quality research to the widest possible audience. Quality We aim for excellence, sustained by a responsible and professional approach to journal publishing. Submissions are guaranteed to receive a decision within 12 weeks. The elapsed time from submission to publication for the articles averages 5-6 months. Indexing of Pertanika Pertanika is now over 30 years old; this accumulated knowledge has resulted in Pertanika journals being indexed in SCOPUS (Elsevier), EBSCO, and AGRICOLA, etc. Future vision We are continuously improving access to our journal archives, content, and research services. We have the drive to realise exciting new horizons that will benefit not only the academic community, but society itself. We also have views on the future of our journals. The emergence of the online medium as the predominant vehicle for the 'consumption' and distribution of much academic research will be the ultimate instrument in the dissemination of research news to our scientists and readers. Aims and scope Pertanika Journal of Tropical Agricultural Science aims to provide a forum for high quality research £ related to tropical agricultural research. Areas relevant to the scope of the journal include: S agricultural biotechnology, biochemistry, biology, ecology, fisheries, forestry, food sciences, genetics, microbiology, pathology and management, physiology, plant and animal sciences, production of plants and animals of economic importance, and veterinary medicine. Editorial Statement Pertanika is the official journal of Universiti Putra Malaysia. The abbreviation for Pertanika Journal of Tropical Agricultural Science is Pertanika J. Trop. Agric. Sci. ARCHIVE COPY Editorial Board (Please Do Not Remove' Editor-in-Chief Tan, S.G., Malaysia I Molecular population genetics Executive Editor Kanwal, Nayan D.S., Malaysia Environmental issues- landscape plant modelling applications Editorial Board Ab-Shukor, N.A. Othman, Rofina Y. Tree improvement, Forestry genetics Agricultural biotechnology & biotechnology Universiti Malaya, Malaysia Universiti Putra Malaysia, Malaysia Radu, S. Ambak, M.A. Food safety. Risk assessment, Fisheries Molecular biology Universiti Malaysia Terengganu, Malaysia Universiti Putra Malaysia, Malaysia Anuar, A.R. Saleh, G.B. Soil fertility and management Plant breeding and genetics Universiti Putra Malaysia, Malaysia Universiti Putra Malaysia, Malaysia Bignell. David E. Salleh, B. Soil biology and termite biology Plant pathologist/Mycologist University of London, U.K Universiti Sains Malaysia, Malaysia Bryden, Wayne L. Saw, L.G. Animal nutrition, Toxicology, Food safety Botany and conservation, Plant ecology University of Queensland, Australia Forest Research Institute Malaysia (FRIM), Kepong, Malaysia Clyde, M.M. Genetics (Cytogenetics) Shamshuddin, J. Universiti Kebangsaan Malaysia, Malaysia Soil science, Soil mineralogy Universiti Putra Malaysia, Malaysia Gan, Yik-Yuen Molecular biology, Genetics, Biotechnology Siddique, K.H.M. Nanyang Technological University, Singapore Crop and environment physiology, Germplasm enhancement Ibrahim, Y.B. The University of Western Australia, Agricultural entomology Australia Universiti Putra Malaysia, Malaysia Tan, W.S. Idris, A. B. Molecular biology. Virology, Protein Entomology (Insect taxonomy and chemistry biodiversity, Integrated pest management, Universiti Putra Malaysia, Malaysia Biological control, Biopesticides) Universiti Kebangsaan Malaysia, Malaysia Yap, C. K. Biology, Ecotoxicology Jamilah, B. Universiti Putra Malaysia, Malaysia Food science and technology, Food quality /processing and preservation Zamri-Saad, M. Universiti Putra Malaysia, Malaysia J cterinaty pathology Universiti Putra Malaysia, Malaysia Editorial Advisory Board Baas, P. Matthews, G. Systematic botany Pest management National Herbarium of The Netherlands, Leiden Imperial College London, U.K University Branch, The Netherlands Napompeth, B. Hughes, Jane M. Entomology Genetics Kasetsart University, Thailand Griffith University, Australia Rahman, A. llyas, Syed M. Plant protection Post harvest engineering and technology' AgResearch, Raukara Research Indian Council of Agricultural Research, Hyderabad, Centre, Hamilton, New Zealand India Walkinshaw, M. Kang, M.S. Biochemistry Plant breeding & genetics University of Edinburgh, Scotland Punjab Agricultural University, India / Louisiana State University Agric. Center Baton Rouge, USA Woodruff, D. Evolution and conservation of Khan, Tanveer N. animal species, International Plant breeding & genetics education Department of Agriculture and Food, South Perth, University of California, San Diego, Western Australia USA Mather, Peter B. Wright, Denis J. Ecology and genetics Pest management Queensland University of Technology, Australia Imperial College London, U.K Editorial Office Pertanika, Research Management Centre (RMC), 4th Floor, Administration Building Universiti Putra Malaysia, 43400 Serdang, Selangor, Malaysia Tel: +603 8946 6192, 8946 6185 • Fax: +603 8947 2075 E-mail: ndeeps(5)admin.upm.edu.my Publisher The UPM Press Universiti Putra Malaysia 43400 UPM, Serdang, Selangor, Malaysia Tel: +603 8946 8855, 8946 8854 • Fax: +603 8941 6172 [email protected] URL: http://penerbit.upm.edu.mv The publisher of Pertanika will not be responsible for the statements made by the authors in any articles published in the journal. Under no circumstances will the publisher of this publication be liable for any loss or damage caused by your reliance on the advice, opinion or information obtained either explicitly or implied through the contents of this publication. All rights of reproduction are reserved in respect of all papers, articles, illustrations, etc., published in Pertanika. All material published in this journal is protected by copyright, which covers exclusive rights to reproduce and distribute the material. No material published in Pertanika may be reproduced or stored on microfilm or in electronic, optical or magnetic form without the written authorization of the Publisher. Copyright © 2009 Universiti Putra Malaysia Press. All Rights Reserved. Pertanika Journal of Tropical Agricultural Science Vol. 32 (2) Aug. 2009 Contents Review Article Net Primary Productivity of Forest Trees: A Review of Current Issue 111 Hazandy Abdul-Hamid, Maurzio Mencuccini and Nor-Aini Ab-Shukor Regular Articles Responses of Musa AAA Berangan to 1-methylcyclopropene 125 Phebe Ding and Khairul Bariah Darduri Juvenile Stem Volume Equations for Planted Azadirachta excelsa in the State of 133 Johore, Peninsular Malaysia Ong Kian Huat, Lim Meng Tsai and John Keen Chubo Biocoversion of Oil Palm Empty Fruit Bunch by Aspergillus niger EB4 under 143 Solid-state Fermentation Azhari Samsu Baharuddin, Nor Asma Abd Razak, Nor Aini Abdul Rahman, Satiawihardja Budiatman, Yoshihito Shirai and Mohd Ali Hassan Selected Articles from the 7th National Genetics Congress 2007 Overexpression of Wildtype Periostin and Transforming Growth Factor Beta I Genes 153 in Colorectal Carcinoma: A Preliminary Study Chia Sze Wooi and Edmund Sim Ui Hang Improved Accuracy for Diagnosis of Nasopharyngeal Carcinoma by the Combination 161 of Recombinant EBV Proteins ZEBRA/IgA and LMP2A/IgG ELISA S.H. Wong, E.L. Tan, C.C. Ng and C.K. Sam Selected Articles from the 3rd Biology Colloquium 2007 New Records of Terrestrial Pteridophytes in Genting Highlands, Pahang, Malaysia 169 Salifah Hasanah Ahmad Bedawi, Rusea Go and Muskhazli Mustafa Antibacterial Activity of Methanolic Crude Extracts from Selected Plant Against 175 Bacillus cereus Muskhazli M., Dirnahayu M., Nor Azwady A.A., Nurhafiza Y., Nor Dalilah E. and Che Ku Nurshaira C.K.N. Selected Articles from the UPM Rice Research Colloquium 2008 Somatic Embryogenesis from Scutellar Embryo of Oryza sativa L. var. MR219 185 Syaiful Bahri Panjaitan, Siti Nor Akmar Abdullah, Maheran Abdul Aziz, Sariah Meon and Othman Omar Influence of Flooding Intensity and Duration on Rice Growth and Yield 195 Abdul Shukor Juraimi, Muhammad Saiful, A.H., Mahfuzah Begum, Anuar, A.R and Azmi, M. Influence of Root Exudate Carbon Compounds of Three Rice Genotypes on 209 Rhizosphere and Endophytic Diazotrophs Naher, U.A, Radziah, O., Halimi, M.S, Shamsuddin, Z.H. and Mohd Razi, I. Upland Rice Varieties in Malaysia: Agronomic and Soil Physico-Chemical 225 Characteristics M.M. Hanafi, A. Hartinie, J. Shukor and T.M.M. Mahmud Seed Germination and Proline Accumulation in Rice (Oryza sativa L.) as Affected by 247 Salt Concentrations Momayezi, M.R., Zaharah, A.R., Hanafi, M.M. and Mohd Razi, I. Upland Rice Root Characteristics and Their Relationship to Nitrogen Uptake 261 Zaharah, A.R. and Hanafi, M.M. Deterministic Model Approaches in Identifying and Quantifying Technological 267 Challenges in Rice Production and Research and in Predicting Population, Rice Production and Consumption in Malaysia Ahmad Selamat and Mohd. Razi Ismail Pollen and Seed Yield Components of Water-stressed Cultivated and Weedy Rice 293 Puteh, A.B, Jali, N., Ismail, M.R., Juraimi, A.S. and Samsudin, N. Critical Period of Weed Competition in Direct Seeded Rice Under Saturated and 305 Flooded Conditions Abdul Shukor Juraimi, M.Y. Mohamad Najib, M. Begum, A.R. Anuar, M. Azmi and A. Puteh Critical Time of Nitrogen Application During Panicle Initiation on the Yield of Two 317 Malaysian Rice Cultivars (Oryza sativa L.) Bah, A., S.R. Syed Omar, A.R. Anuar and M.H.A.Husni Growth, Physiological and Biochemical Responses of Malaysia Rice Cultivars to 323 Water Stress Wan Mohammad Zulkarnain, Mohd Razi Ismail, M. Ashrafuzzaman, Halimi Mohd Saud and Ismail C. Haroun Biochemical Diversity of Bacterial Isolates from Paddy Soils of Peninsular Malaysia 335 Maszlin Mohd. Yusof, Halimi Mohd. Saud and Tan My Pein ISSN: 1511-3701 Pertanika J. Trop. Agric. Sci. 32 (2): 111 - 123 (2009) © Universiti Putra Malaysia Press Review Article Net Primary Productivity of Forest Trees: A Review of Current Issue Hazandy Abdul-Hamid1,2*, Maurizio Mencuccini3 and Nor-Aini Ab-Shukor1,2 1Institute of Tropical Forestry and Forest Products, Universiti Putra Malaysia, 43400 UPM, Serdang, Selangor, Malaysia 2Faculty of Forestry, Universiti Putra Malaysia, 43400 UPM, Serdang, Selangor, Malaysia 3School of GeoSciences, University of Edinburgh, EH9 3JU, UK *E-mail: [email protected] ABSTRACT Forest growth is important both economically and ecologically and it follows a predictable general trend with age. Generally, the growth of all forests accelerates as canopies develop in young forests and declines substantially soon after the maximum leaf area is attained. The causes of this decline trend are multiple. Initially, age- and size-related declines were attributed to photosynthesis-respiration imbalance. Subsequently, several competing hypotheses have been proposed over the years, although nutrient and hydraulic limitation hypotheses appear to be the most likely to have caused it. In this paper, the authors attempt to review these hypotheses and concentrate on one related aspect, as this can set the scene for further examining the issues of age-related versus size-related processes. Keywords: Forest growth, canopies, photosynthesis – respiration imbalance GENERAL BACKGROUND fluxes in forest ecosystems and their influence on the global CO cycles. The rate of production of the biomass by both 2 Despite many studies attempting to explain individual trees and forest stands decreases the age-related decline of forest productivity substantially with age (Assmann, 1970). In through a number of theories, the age-related particular, age-related reduction in biomass regulation still remains unclear. There is accumulation is an important consideration in little direct evidence which explains that old the mechanistic models which predict the forest forests assimilate less carbon than the young growth and determine the capacity of the forests forests. This makes model-based predictions to act as carbon sinks. With the current impetus of the carbon sequestration suspect, as most towards silvicultural approaches that employ physiological process models predict carbon uneven-aged systems and extended rotations, assimilation were based on the measurements understanding the physiological basis for the of photosynthesis on young trees (Ryan et al., age-related decline in productivity of dominant 1997a). For example, Bond (2000) provided a species has become increasingly important. list of studies exhibiting two contrasting results Understanding this component of productivity is in the changes in the net photosynthesis with age crucial to quantifying and manipulating carbon of trees and shrubs (Table 1). Received: 4 March 2008 Accepted: 5 May 2009 *Corresponding Author Hazandy Abdul-Hamid, Maurizio Mencuccini and Nor-Aini Ab-Shukor TABLE 1 Studies reporting a comparison of the net photosynthesis in differently aged trees and shrubs Decreased net photosynthesis (A ) with increased ages of trees and shrubs net Species Comparison Reference Chrysothamnus nauseosus Maximum A in summer. Juvenile versus Donovan and Ehleringer net mature. (1992)* Juniperus occidentalis Juvenile versus young mature. Miller et al. (1995)* Larrea tridentate Daily maximum A , three summer months. Franco et al. (1994)* net Juvenile versus mature. Picea abies Light-saturated A , open-grown trees. Kull and Koppel (1987)* net Juvenile versus mature. Picea rubens Seasonal A . Mature versus relatively old- Day et al. (2001) net growth. Pinus aristata Light-saturated A , optimal conditions. Schoettle (1994)* net Mature versus old-growth. Pinus contorta Light-saturated A . Mature versus old- Yoder et al. (1994)* net growth. Pinus ponderosa Light-saturated A . Mature versus old- Yoder et al. (1994)* net growth. Pinus ponderosa Mean morning A . Juvenile versus mature Kolb and Stone (2000)* net and old-growth. Prosopis glandulosa Daily maximum A , after rainfall. Juvenile DeSoyza et al. (1996)* net versus mature. Prunus serotina Growing season average A . Seedling versus Fredericksen et al. (1996)* net sapling versus mature. Sequoiadendron giganteum Maximum A . Seedlings versus juvenile, Grulke and Miller (1994)* net mature and old-growth. No difference or increased net photosynthesis with increased ages of trees and shrubs Species Comparison Reference Acer negundo Maximum A , during peak of drought period. Donovan and Ehleringer net Juvenile versus mature. (1992)* Artemesia tridentata Maximum A in mid-summer at a dry site. Donovan and Ehleringer net, Juvenile versus mature. (1992)* Chrysothamnus nauseosus Maximum A , in mid-summer at a dry site. Donovan and Ehleringer net Juvenile versus adult. (1992)* Prosopis glandulosa Daily maximum A . Summer drought. Small DeSoyza et al. (1996)* net (0.5 m) versus large (1.1 m). Pseudotsuga menziesii No clear trend of light-saturated A . Mature McDowell et al. (2002) net versus old growth. Quercus rubra Light-saturated A , during growing season. Hanson et al. (1994)* net Seedling versus mature. Note: * = cited from Bond (2000). 112 Pertanika J. Trop. Agric. Sci. Vol. 32 (2) 2009 Net Primary Productivity of Forest Trees: A Review of Current Issue The causes of this age-related decline are from juvenile and mature trees, ranging from 1 likely multiple, but the most important potential to 74 years of age. causes which can explain a decline in forest Mencuccini et al. (2005) also used propagated productivity with stand development are nutrient material to separate the relative effects of size limitation, hydraulic limitation and maturation and age on tree growth and metabolism. These (genetic programming) hypotheses. According authors selected four different tree species. to some authors, the so called “age-related” Three of them (one conifer, Scots pine and decline in the forest productivity is primarily two angiosperms with diffuse-porous wood, a “size-related” decline (Weiner and Thomas, sycamore and ring-porous wood, ash) were 2001). Ideally, a test should be conducted on propagated by grafting, whereas the fourth the trees of similar size, but different ages or (poplar) was propagated by direct rooting. vice versa; thereby separating the confounded Poplar was also chosen because of its clonal factors of size and age. An alternative to obtain nature, which allowed a different approach in the trees with similar size but with different ages is separation of age versus size (cf., Mencuccini, to graft scions of the different ages onto young 2007 for further details). The approach adopted rootstocks. in Mencuccini et al. (2005) avoided some of the Studies using grafting have been conducted potential artefacts presented in the earlier works by a few researchers (summarised in Table 2) (Table 2). For instance, the individuals measured in relation to the effect of age on the growth, in the field were almost always the exact same morphology and physiology of grafted scions. donors from where the grafted twigs were taken, However, these studies still yielded contrasting thereby avoiding genetic differences between outcomes with the age of scion. Despite the the donor trees and the grafted plants. For the presence of these grafting studies, there are some conifer in the study by Mencuccini et al. (2005) weaknesses which have never been tackled by (Scots pine), the measurements took place five the previous researchers. For instance, no study to seven years after the initial grafting, to ensure has so far simultaneously compared individuals that enough time had elapsed since grafting had in the field and genetically identical grafted taken place. More importantly perhaps, the seedlings. Day et al. (2001) conducted a similar grafting technique did not leave any room for study, but the individuals selected in the field competition between shoots of the grafted plant were not genetically identical with the grafted and the shoots of the rootstock, as the canopy of seedlings (although they came from the same the accepting rootstock was completely pruned population). Moreover, most of the published away after grafting. evidences on grafting have been obtained from Therefore, additional comparative studies very young trees. In relation to the studies are required to determine whether the different presented in Table 2, for instance, Hutchison et conclusions reached by some of the earlier al. (1990) used individuals taken from four age studies were the results of different species- classes, ranging from 1 to 45 years; whereas, specific developmental events or whether they Rebbeck et al. (1993) used only juvenile and are determined by the different experimental mature (>50 years old) scions. In addition, protocols. Greenwood (1984) studied shoot development as a function of age on Picea taeda, but the AGE- AND SIZE-RELATED comparisons were only made among scions up REGULATION OF THE NET PRIMARY to 12 years of age. Furthermore, Greenwood PRODUCTION et al. (1989) also used grafting approach to study the effect of age on the morphological Observations have showed that aboveground net characteristics and DNA methylation of Larix primary productivity negatively correlates with laricina, but the scions involved were taken the age of tree for both individual trees and single Pertanika J. Trop. Agric. Sci. Vol. 32 (2) 2009 113 Hazandy Abdul-Hamid, Maurizio Mencuccini and Nor-Aini Ab-Shukor TABLE 2 Summary of the results showing contrasting trends involving in grafted scions Species Results Reference Hedera helix Increased light-saturated A with increased ages Bauer and Bauer (1980) net of scions. Juvenile versus mature scions grafted to juvenile rootstock. Larix laricina Height and diameter growth decreased with Greenwood et al. (1989) increased ages of scions. Juvenile versus mature scions grafted to juvenile rootstock. Larix laricina Total chlorophyll increased with increased ages Greenwood et al. (1989) of scions. Juvenile versus mature scions grafted to juvenile rootstock. Larix laricina Increased light-saturated A with increased ages Hutchison et al. (1990) net (indoor-grown trees) of scions. Juvenile versus mature scions grafted to juvenile rootstock. Larix laricina No trend observed in light-saturated A with Hutchison et al. (1990) net (outdoor-grown trees) increased ages of scions juvenile versus mature scions grafted to juvenile rootstock. Larix laricina Increased xylem diameters with increased ages Takemoto and Greenwood of scions. Juvenile versus mature scions grafted (1993) to juvenile rootstock. Picea rubens Decreased A with increased ages of scions. Rebbeck et al. (1993) net Juvenile versus mature scions grafted to juvenile rootstock. Picea rubens Decreased light-saturated A with increased age Day et al. (2001) net of scions. Mature versus old scions. Picea taeda Diameter and height growth, leaf area, branch Greenwood (1984) numbers and scion biomass decreased with increased scion ages. Juvenile versus young scions grafted to juvenile rootstock. Pinus radiata Diameter and height growth and branch numbers Sweet (1973) decreased with increased scion ages. Juvenile versus mature scions. Pseudotsuga menziesii Diameter and height growth, branch numbers Ritchie and Keeley (1994) and branch length decreased with increased scion ages. Juvenile versus young scions grafted to juvenile rootstock. cohort stands (Assmann, 1970; Whittaker, 1975; and ponderosa pine may be a significant cause Bormann and Likens, 1979; Harcombe et al., of net production decreases in old trees. This 1990). Long-term studies on even-aged forests is supported by several studies conducted on showed that the maximum bole increment rate age-related differences in photosynthetic rates in occurs shortly after crown closure and declines Scots pine (Kull and Koppel, 1987), bristlecone as trees reach maturity (Assmann, 1970). pines (Schoettle, 1994) and hybrid Englemann Yoder et al. (1994) suggested that age-related x white x Sitka spruce (Richardson et al., 2000). declines in photosynthetic rates for lodgepole However, photosynthetic rates in the eastern 114 Pertanika J. Trop. Agric. Sci. Vol. 32 (2) 2009
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