A functional-structural model for radiata pine (Pinus radiata) focusing on tree architecture and wood quality

Backgrounds and Aims

Functional–structural models are interesting tools to relate environmental and management conditions with forest growth. Their three-dimensional images can reveal important characteristics of wood used for industrial products. Like virtual laboratories, they can be used to evaluate relationships among species, sites and management, and to support silvicultural design and decision processes. Our aim was to develop a functional–structural model for radiata pine (Pinus radiata) given its economic importance in many countries.

Methods

The plant model uses the L-system language. The structure of the model is based on operational units, which obey particular rules, and execute photosynthesis, respiration and morphogenesis, according to their particular characteristics. Plant allometry is adhered to so that harmonic growth and plant development are achieved. Environmental signals for morphogenesis are used. Dynamic turnover guides the normal evolution of the tree. Monthly steps allow for detailed information of wood characteristics. The model is independent of traditional forest inventory relationships and is conceived as a mechanistic model. For model parameterization, three databases which generated new information relating to P. radiata were analysed and incorporated.

Key Results

Simulations under different and contrasting environmental and management conditions were run and statistically tested. The model was validated against forest inventory data for the same sites and times and against true crown architectural data. The performance of the model for 6-year-old trees was encouraging. Total height, diameter and lengths of growth units were adequately estimated. Branch diameters were slightly overestimated. Wood density values were not satisfactory, but the cyclical pattern and increase of growth rings were reasonably well modelled.

Conclusions

The model was able to reproduce the development and growth of the species based on mechanistic formulations. It may be valuable in assessing stand behaviour under different environmental and management conditions, assisting in decision-making with regard to management, and as a research tool to formulate hypothesis regarding forest tree growth and development.     

Quantification of compression wood severity in tracheids of Pinus radiata D. Don using confocal fluorescence imaging and spectral deconvolution

Confocal fluorescence microscopy was used to examine the spectral characteristics of lignin autofluorescence in secondary cell walls of normal and compression wood from Pinus radiata. Using UV excitation, fluorescence spectra of normal and compression wood sections showed significant differences, especially in the outer secondary cell wall of tracheids, with a shift in maxima from violet to blue wavelengths between normal and compression wood. A comparison of normal wood, mild and severe compression wood, showed that the wavelength shift was intermediate in the mild compression wood compared to the severe compression wood, thus offering the possibility of quantifying the severity by measuring ratios of fluorescence at violet and blue wavelengths. Fluorescence induced by blue light, rather than UV, was less well differentiated amongst wood types. Spectral deconvolution indicated the presence of a minimum of five discrete lignin fluorophores in the cell walls of both normal and compression wood tracheids. Comparison with lignin model compounds suggest that the wavelength shift may correspond in part to increased levels of p-hydroxy type lignin in the compression wood samples. The combination of confocal fluorescence imaging and related spectral deconvolution therefore offers a novel technique for characterising cell wall lignin in situ.     

Modelling environmental variation in Young's modulus for Pinus radiata and implications for determination of critical buckling height

• Background and Aims Although density-specific stiffness,E/, (where E is Young's modulus and is wood density) is oftenassumed constant by the elastic similarity model, and in determinationof critical buckling height (H_crit), few studies have testedthis assumption within species. Here this assumption is testedfor Pinus radiata growing across an environmental gradient,and theory is combined with data to develop a model of Young'smodulus. • Methods Analyses use an extensive series of environmentalplots covering the range of climatic and edaphic conditionsover which P. radiata is grown in New Zealand. Reduced majoraxis regression was used to determine scaling exponents betweenlog–log plots of H_crit vs. groundline diameter (D), andE/ vs. D. Path analysis was used to identify significant directand indirect (through stem slenderness) edaphic and climaticinfluences on E. • Key Results Density-specific stiffness exhibited 3-foldvariation. As E/ scaled positively with D, the exponent of 0·95between H_crit and D exceeded the assumed value of 0·67under constant E/. The final path analysis model included meanair temperature in early autumn (T_aut) and slenderness as significant(P < 0·05) positive direct influences on E. Tree leafarea index and T_aut were indirectly associated with E throughtheir significant (P < 0·05) positive direct relationshipwith stem slenderness. Young's modulus was most sensitive toT_aut, followed by stem slenderness then leaf area index, andthe final model explained 76 % of the variance in E. • Conclusions The findings suggest that within speciesE/ variation may influence H_crit and the scaling exponent betweenD and H_crit so important in assumptions regarding allometricrelationships. The model presented may provide a useful meansof determining variation in E, E/ and H_crit across environmentalgradients.     

Photosynthesis and wood structure in Pinus radiata D. Don during dehydration and immediately after rewatering

Abstract. Experiments were carried out on Pinus radiata (D. Don) trees grown as cuttings from clonal parent stock. Some of these trees were about 0.4 m high while others were about 5 m high; all were grown in containers. The stem diameters at the tops and at the bottoms of the large trees, rates of photosynthesis, and needle water potentials were measured both when the trees were well watered and as they dehydrated after water was withheld. The water potentials of well-watered plants was highest in the small trees and lowest at the top of the large trees. When water was withheld, photosynthesis was in most cases unaffected by a small reduction in water potential, but the rate of photosynthesis fell as water potentials declined further. The stems of the large trees expanded at a constant rate when the trees were well watered and for part of the dehydration period, while subsequent stem shrinkage and the fall in photosynthesis both occurred at approximately the same time.
Water potentials increased little in the 24 h after rewatering, and significant rates of photosynthesis were not measured until 2 or 3 d later while renewed stem expansion was not measured until 2 d after rewatering.
Water deficits reduced the lumen diameter of newly matured stem tracheids, but increased the thickness of their walls. After 1 month of water potentials of about −2.4 MPa, tracheid lumen diameter and wall thickness were both much reduced, and this reduction continued in tracheids maturing shortly after rewatering.     

Genetic control of growth,biomass allocation,and survival under drought stress in Pinus radiata D. Don seedlings

Drought is one of the critical factors limiting plant growth, and knowledge about genetic adaptation to drought is necessary to develop strategies for successful reforestation on drought-prone sites. In this study, genetic variation was investigated in 98 full- and half-sib radiata pine (Pinus radiata D. Don) families, representing Chilean coastal and interior populations, subjected to two water regimes: well-watered and drought-stressed. Assessed traits, in 5-month-old seedlings, included height (H), diameter (D), height-to-diameter ratio (HDR), dry biomass of needles (NDW), stem (SDW), roots (RDW), and total (TDW), root-to-shoot ratio (RSR), and survival (SUR). After 115 days of treatment, growth, biomass, and survival were nearly two times higher under the well-watered regime than under the stressed one. Families differed significantly in most traits, with individual tree heritabilities ranging from 0.14 for SUR to 0.63 for D in the well-watered treatment. Families from the interior showed the highest heritability for D, SDW, RDW, and TDW when grown in the water-stress treatment. The genetic correlations between treatments were moderately strong, which suggests the presence of a genotype by watering regime interaction. Most traits were strongly correlated (genetic correlations often exceeded 0.40). Compared to the first generation families from coastal sites, the third generation families from the interior sites showed an increase in SUR and RSR. Thus, potential exists to screen families at the seedling stage for drought hardiness and to identify parents from the interior sites with potential to produce a more drought-resistant breed with satisfactory growth rates and yields in dry environments.     

Molecular control of early cone development inPinus radiata

Summary A family of genes expressed during early stages of shoot development were isolated fromPinus radiata. A homologue of theLEAFY/FLORICAULA flower meristem-identity genes,NEEDLY (NLY), and three MADS-box genes,PrMADS1, PrMADS2 andPrMADS3 (Pinus radiata MADS-box genes), were expressed at early stages of initiation and differentiation of reproductive (male and female) cone buds, as well as vegetative buds. Expression ofNLY in transgenicArabidopsis thaliana promoted floral fate, demonstrating that it encodes a functional ortholog of theFLORICAUL A/LEAFY genes of angiosperms.Abbreviations DSB dwarf shoot bud - LSTB long-shoot terminal bud - PCB pollen cone bud - SCB seed cone bud - LD long day - SD short day     

Characterization of white zones produced on Pinus radiata wood chips by Ganoderma australe and Ceriporiopsis subvermispora

White zones produced on biodegraded Pinus radiata wood chips were characterized by micro-localized-FTIR (Fourier Transformed Infra Red) spectroscopy and scanning electron microscopy. Both techniques permitted assignment of the white zones to a selective lignin removal process. Although both fungi studied have degraded lignin selectively in these restricted superficial areas, chemical analysis of the wood chips indicated that Ganoderma australe removed 16% of the initial amount of glucan at the 20% weight loss level. Ceriporiopsis subvermispora did not remove glucan at weight loss values below 17%. Prolonged biodegradation resulted in reduction of white zones by G. australe, and increased white zones from C. subvermispora decayed samples. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photoperiod control of the initial phase of frost hardiness development in Pinus radiata

Abstract. The development of frost hardiness in Pinus radiata was investigated to establish whether there is quantitative relationship between photoperiod and the hardening process. Three controlled environment experiments were carried out. In the first, seedlings were exposed to a photoperiod that shortened from 13 h at a rate of 3 min d^?1 to 9.5 h. At intervals, the photoperiod was either held constant or lengthened. In the second experiment, seedlings were exposed to one of five constant photoperiods between 9 and 12 h for up to 90 d. In the third, seedlings were exposed to photoperiods shortening at rates of 1 or 5 min d^?1. Frost hardiness was also measured during the natural photoperiod-controlled stage of hardening in outdoor-grown seedlings. Frost hardiness developed at a constant rate in response to a shortening photoperiod once it had declined to about 12 h. This rate was consistent with the hardening rate that occurred in outdoor-grown seedlings. Hardening stopped when the photoperiod became constant, indicating a tight coupling between changes in photoperiod and hardiness development. When the photoperiod was held constant, the extent of frost hardiness was directly dependent on the photoperiod but the rate of hardening was apparently independent of the length of photoperiod. However, the rate of hardening was dependent on the rate at which the photoperiod shortened, increasing linearly with increases in the rate of change in photoperiod between 0 and 3 min d^?1. These results suggest shortening photoperiods control the first stage of the hardening process by regulating the rate of hardening. Frost hardening was inherently unstable once the maximum hardiness was reached since spontaneous dehardening occurred in spite of the controlled conditions. Dehardening also occurred when the photoperiod was lengthened suggesting that the cue for dehardening to commence was the shift from shortening to lengthening photoperiods.     

A new method for vulnerability analysis of small xylem areas reveals that compression wood of Norway spruce has lower hydraulic safety than opposite wood

Compression wood is formed at the underside of conifer twigs to keep branches at their equilibrium position. It differs from opposite wood anatomically and subsequently in its mechanical and hydraulic properties. The specific hydraulic conductivity (k_s) and vulnerability to drought‐induced embolism (loss of conductivity versus water potential ψ) in twigs of Norway spruce [Picea abies (L.) Karst.] were studied via cryo‐scanning electron microscope observations, dye experiments and a newly developed ‘Micro‐Sperry’ apparatus. This new technique enabled conductivity measurements in small xylem areas by insertion of syringe cannulas into twig samples. The hydraulic properties were related to anatomical parameters (tracheid diameter, wall thickness). Compression wood exhibited 79% lower k_s than opposite wood corresponding to smaller tracheid diameters. Vulnerability was higher in compression wood despite its narrower tracheids and thicker cell walls. The P₅₀ (ψ at 50% loss of conductivity) was ?3.6 MPa in opposite but only ?3.2 MPa in compression wood. Low hydraulic efficiency and low hydraulic safety indicate that compression wood has primarily a mechanical function.     

Stand variation in Pinus radiata and its relationship with allometric scaling and critical buckling height

Background and Aims

Allometric relationships and the determination of critical buckling heights have been examined for Pinus radiata in the past. However, how they relate to more mature Pinus radiata exhibiting a wide range of stem diameters, slenderness and modulus of elasticity (E) at operationally used stand densities is largely unknown. The aim of this study was to examine the relationship between Pinus radiata stand structure variables and allometric scaling and critical buckling height.

Methods

Utilizing a Pinus radiata Nelder trial with stand density and genetic breed as variables, critical buckling height was calculated whilst reduced major axis regression was used to determine scaling exponents between critical height (H_crit), actual height (H), ground line diameter (D), slenderness (S), density-specific stiffness (E/ρ) and modulus of elasticity (E).

Key Results

Critical buckling height was highly responsive to decreasing diameter and increasing slenderness. Safety factors in this study were typically considerably lower than previously reported margins in other species. As density-specific stiffness scaled negatively with diameter, the exponent of 0·55 between critical height and diameter did not meet the assumed value of 0·67 under constant density-specific stiffness. E scaled positively with stem slenderness to the power of 0·78.

Conclusions

The findings suggest that within species density-specific stiffness variation may influence critical height and the scaling exponent between critical height and diameter, which is considered so important in assumptions regarding allometric relationships.     

Isolation of Pinus radiata Genomic DNA Suitable for RAPD Analysis

A protocol is presented for Pinus radiata genomic DNA isolation based on an alkyltrimethyl-ammonium bromide (CTAB) method described for other woody species. The method involves mortar grinding of tissue, a modified CTAB extraction employing high salt concentrations and polyvinyl pyrrolidone, a RNase A treatment and successive isoamyl alco- hol-chloroform extractions. The yield was approximately 15 g DNA per 100 mg of initial fresh plant material. The genomic DNA obtained by this method was suitable to be used in simple sequence repeat and random-amplified-polymorphic DNA reactions. This extraction method would allow the molecular analysis of shoots from different clones within P. radiata families.     

A diffusible auxin from Pinus radiata pollen and its possible role in stimulating ovule development

Summary Germinated pollen of Pinus radiata contains an auxin which is active in the Avena coleoptile test. It differs from all other hormones detected in pine pollen in that it is readily able to diffuse out from the pollen into an agar medium. It is suggested that, following pollination in vivo, the auxin may diffuse from the germinated pollen-tube onto the nucellus, thereby triggering the processes which allow ovule and gametophyte development to proceed. The auxin is water-soluble and may be an indole derivative.     

Fine Structure of Parenchymatous and Differentiated Pinus radiata Callus

The fine structure of Pinus radiata D. Don callus before andafter differentiation into stem-like tissues has been examinedwith the electron microscope. In callus prior to differentiation(here called parenchymatous callus) the cells accumulate tanninsas they age and are quite distinct from the cells of differentiatedcallus. In the latter, cambium, phloem and xylem cells may beidentified by their general morphology and by their ultrastructuralfeatures. Differentiation into a true stem-like structure is,however, incomplete in that the tissues are not uniformly oriented,and parenchyma cells of the rays and phloem contain chloroplasts.The tracheids also show unusual differentiation in that borderedpits form over their entire surface and may be of two types.The reasons for these variations are discussed.     

Climatic control of intra-annual wood density fluctuations of Pinus pinaster in NW Spain

Tree rings of Pinus pinaster often contain intra-annual density fluctuations (IADFs), which have been attributed to the succession of dry and rainy periods typical of Mediterranean climate, but their formation has not been studied yet under Atlantic climate. We analyzed the occurrence and climatic significance of replicated IADFs in ten monospecific stands in NW Spain. The frequency of IADFs was higher than previously reported for this species under Mediterranean conditions and consistently decreased with increasing elevation. The formation of bands of latewood-like tracheids within the earlywood was favored by dry previous August, cold previous winter and dry April. Bands of earlywood-like tracheids within the early latewood were also favored by low winter temperatures. However, their occurrence was geographically heterogeneous, with two groups of stands being defined by their distances to the shoreline. In coastal stands, cold May–August triggered IADFs formation, while in inland stands their formation was favored by dry May–July. Regional winter temperatures and April water balance were strongly related to the East Atlantic (EA) pattern, which greatly conditioned the occurrence of IADFs in the earlywood and the early latewood. By contrast, the presence of bands of earlywood-like tracheids in the late latewood was independent of the EA pattern, being strongly related to warm conditions in spring and especially to a wet October. The link between regional climate and the EA pattern strongly controlled the physiological processes that determine intra-annual growth dynamics and short-term cell enlargement of P. pinaster in NW Spain.     

The origin and genetic diversity of Pinus radiata in Australia

Summary Despite the fact that forest trees are in early stages of domestication there has been little direct evaluation of either the origin of, or genetic diversity within the breeding material in tree improvement programs. Allozyme variation was used to compare the total genetic diversity in the breeding programs of P. radiata within Australia and the five wild populations in North America. The current breeding populations were very similar genetically and were essentially homogenous with only 1.8% of the variation among programs. The total genetic diversity in the species was 0.12, which is a low estimate compared to most conifers. Overall in the Australian material the genetic diversity was somewhat less. The comparison of allelic frequencies in the five native populations with the Australian material indicates that the Monterey and Año Nuevo populations were probably the major source of the original introductions and that a substantial portion of the genetic diversity in the two populations has been captured in current breeding programs. The three southern populations do not appear to be currently represented in the breeding programs. The implications for future breeding strategies are discussed.     

Evaluation of AFLP for genetic mapping in Pinus radiata D. Don

Efficient construction of reasonable density genetic linkage maps is an essential component of QTL detection programmes. The AFLP technique has been used to produce genetic linkage maps in a range of species. We have developed protocols to generate reproducible AFLP profiles in Pinus radiata and have evaluated the inheritance and informativeness of AFLP markers in this important timber species. The large genome size of P. radiata necessitated increased levels of selection at both the pre-amplification and selective amplification steps of the AFLP protocol to generate reproducible AFLP profiles. Once optimised ca. 41.3 scorable AFLP bands were resolvable through denaturing gels, of which 48.4% were polymorphic in a screen of eight unrelated trees. This level of polymorphism is ca. three times higher than with RAPD markers. The total number of bands and the number of polymorphismic bands per PCR were ca. halved when AFLPs were electrophoresed on non-denaturing gels and stained with ethidium bromide. Using the protocols developed, AFLP is an efficient method for generating the DNA markers required for genetic linkage map construction in P. radiata. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mild hydrogenolysis of in-situ and isolated Pinus radiata lignins

The Pd/C-catalysed hydrogenolysis of in-situ and isolated lignins from Pinus radiata wood was investigated to gain a more complete understanding of the factors affecting yield and composition of the hydrogenolysis products. Such hydrogenolysis products could potentially be refined into aromatic feedstock chemicals providing sustainable alternatives to petroleum-derived phenols. Lignins were converted into solvent-soluble oils composed of monomeric, dimeric and oligomeric products in high yields, up to 89% of the original lignin. The main monomer products were dihydroconiferyl alcohol and 4-n-propyl guaiacol. Dimeric and oligomeric compounds constituted 75% of the hydrogenolysis oils and were mainly composed of dihydroconiferyl alcohol and 4-n-propyl guaiacol units linked by β-5, 5-5, 4-O-5 and β-1 linkages. Hydrogenolysis of steam exploded wood gave lower yields of lignin hydrogenolysis products compared to unmodified wood due to fewer reactive aryl-ether linkages in the lignin.