RADIATA PINE Bc. Radim Lainka

TFA Forest dendrology, genetics and tree breeding SS 2015

COMMON NAMES

Radiata Pine

Monterey Pine

Pin insigne

Pin géant

Pino insigne

Pinheiro insigne

Pino de Monterey

insignis pine

pino quebradizo Fig. 1 Guadalupe Island pines(LITTLE, ELBERT L. Jr.. 1980. The Audubon Society field guide to North American trees. New York: Alfred A. Knopf.,

ÚRADNÍČEK, BROUMOVSKÝ and UNČOVSKÝ,1990. Lesní hospodářství v tropech a subtropech-Vybrané dřeviny, MZLU http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G59615.html)

(http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=183372) PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands.(http://www.prota4u.org/protav8.asp?en=1&p=Pinus+radiata+D.Don)

TAXONOMIC HIERARCHY

Kingdom Plantae – plantes, Planta, Vegetal, plants

Subkingdom Viridiplantae

Infrakingdom Streptophyta – land plants

Superdivision Embryophyta

Division Tracheophyta – vascular plants, tracheophytes

Subdivision Spermatophytina – spermatophytes, seed plants, phanérogames

Class Pinopsida – conifers

Subclass Pinidae

Order Pinales – pines

Family Pinaceae – pines

Genus Pinus L. – pine

Species Pinus radiata D. Don –> var. radiata/var. pinata

Retrieved [March, 5, 2016], from the Integrated Taxonomic Information System (ITIS) on-line database (http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=183372) The Gymnosperm database (http://www.conifers.org/pi/Pinus_radiata.php)

PLANT CHARACTERISTICS

H. 15-30 (64) m,

DBH 30-90 (280) cm

contorted to straight, crown broadly conic

twigs - sometimes glaucous, aging gray, rough, slender, red-brown,

buds - resinous, ovoid to ovoid-cylindric, red-brown, ca. 1.5 cm

needles - 2 (var. binata) or 3 (type variety) per fascicle, spreading-ascending, persisting 3-4 years, (8)9-15(20) cm × 1.3-1.8(2) mm, straight, slightly twisted, deep yellow-green

(LITTLE, ELBERT L. Jr.. 1980. The Audubon Society field guide to North American trees. New York: Alfred A. Knopf. KRAL, R. 1993. Pinus. Flora of North America Editorial Committee (eds.): Flora of North America North of Mexico, Vol. 2. Oxford University Press.

M.P. Frankis e-mail 1999.03.05, pers. obs.)

Fig. 2 Guadalupe Island pine trunk

Fig. 3 Guadalupe Island pine bark

ko

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G61056.html)

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G58616.html)

PLANT CHARACTERISTICS

Pollen cones: ellipsoid-cylindric, 10-15 mm, orange-brown

Seed cones: maturing in February, 2 years after pollination, persistent 6-20(40) years, often serotinous, numerous, solitary to whorled, 7-15 cm, yellow-brown, lustrous, scales rigid, stalks to 1 cm

Seeds: body ca. 6 mm, dark brown, wing 20-30 mm, compressed-ellipsoid

(LITTLE, ELBERT L. Jr.. 1980. The Audubon Society field guide to North American trees. New York: Alfred A. Knopf. KRAL, R. 1993. Pinus. Flora of North America Editorial Committee (eds.): Flora of North America North of Mexico, Vol. 2. Oxford University Press.

M.P. Frankis e-mail 1999.03.05, pers. obs.)

Fig. 4 Close-up of Guadalupe Island pine needles

Fig. 5 Immature cone of Guadalupe Island pine

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G61055.html)

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G58620.html)

Fig. 6 Guadalupe Island pine cones

Fig. 7 Male Guadalupe Island pine flowers Microsporangiate Strobili with bloom period

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G59622.html)(http://www.pinetum.org/Lovett/pinecones.htm)

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G91136.html)Calflora: Information on California plants for education, research and conservation, with data contributed by public and private institutions and individuals, including the Consortium of California Herbaria. [web application]. 2016. Berkeley, California: The Calflora Database [a non-profit organization]. Available: http://www.calflora.org/ (Accessed: Mar 05, 2016).

ORIGIN AND DISTRIBUTION

Naturally - three localities in a fog belt on the coast of central California (at 30-400 m elevation; San Mateo and Santa Cruz counties, Monterey County, and in San Luis Obispo County)

var. binata - Islas Guadalupe and Cedros, off the west coast of Baja California Norte, Mexico (at 600-1200 m elevation)

Timber tree in vast areas of New Zealand (the most common tree), Australia, Chile, Argentina, Uruguay, SW Europe, Kenya, Ghana, Nigeria, Sudan, Ethiopia, Tanzania, Malawi, Zimbabwe, South Africa and Madagascar.

(ROGERS, D. L., 2002. In situ genetic conservation of Monterey pine (Pinus radiata D. Don): Informations and recommendations. Genetic Resources Conservation Program, Division of Agriculture and Natural Resources, University of California, Davis, C. A..

LITTLE, ELBERT L. Jr.. 1980. The Audubon Society field guide to North American trees. New York: Alfred A. Knopf. KRAL, R. 1993. Pinus. Flora of North America Editorial Committee (eds.): Flora of North America North of Mexico, Vol. 2. Oxford University Press.

Richardson, D.M. (ed.). 1998. Ecology and Biogeography of Pinus. Cambridge University Press. ISBN 0-521-55176-5. http://www.iucnredlist.org/details/42408/0)

Fig. 8a/b Natural distribution of the type variety of Pinus radiata

GRIFFIN, J.R. and W.B. CRITCHFIELD. 1972. The distribution of forest trees in California. Berkeley: U.S.D.A. Forest Service. http://www.na.fs.fed.us/spfo/pubs/silvics_manual/Volume_1/pinus/radiata.jpg) LEDIG, F.T., J.J. VARGAS HERNÁNDEZ, & K.H. JOHNSES. 1998. The Conservation of Forest Genetic

Resources - Case Histories from Canada, Mexico, and the United States. Journal of Forestry, 96(1): 32-41.(http://www.fwpa.com.au/images/processing/PNC135-0809_Native_radiata_germplasm_conservation_Research_Report_0.pdf)

Fig. 8b Adapted from LEDIG et al. (1998)

Fig. 8a Adapted from GRIFFIN, J.R. and W.B. CRITCHFIELD ( 1972)

Pinus radiata var. radiata

Pinus radiata var. binata

Fig. 9 Actual Distribution of Radiata pine in California (Caflora)

Calflora: Information on California plants for education, research and conservation, with data contributed by public and private institutions and individuals, including the Consortium of California Herbaria. [web application]. 2016. Berkeley, California: The Calflora Database [a non-profit organization]. Available: http://www.calflora.org/ (Accessed: Mar 05, 2016).

Fig. 10 Point map of Radiata pine – world scale

(http://www.discoverlife.org/mp/20m?map=Pinus+radiata)

ASSOCIATED ORGANISMS

beneficials (not specific):

butterflies

Pine White Nephasia menapia

pests (not specific):

leafhoppers

Glassy-winged sharpshooter Homalodisca vitripennisCalflora: Information on California plants for education, research and conservation, with data contributed by public and private institutions and individuals, including the Consortium of California Herbaria. [web application]. 2016. Berkeley, California: The Calflora Database [a non-profit organization]. Available:https://www.calflora.org/entry/plantchar.html?crn=6523

ENVIRONMENTAL REQUIREMENTS AND ECOLOGY

Tab. 1 Characteristics of five native radiata pine populations

Population Latitude (°N)

Altitude (m)

Raiffall aprox. (mm)

Area (ha) Soil

Año Nuevo 37 10-30 800 450 Fine loams, depth variable

Monterey 36.5 10-40 400 3800 Very varied fertility and base status

Cambria 35.5 10-00 500 900Sandy loam,

localised poor drainage

Guadalupe Island 29 400-1200 300? 220 trees Rocky loam

Cedros Island 28 380-640 200? 150 Skeletal

(ELRIDGE, K.G. 1978. Refreshing the genetic resources of radiata pine plantations. CSIRO Division of Forest Research, Genetics Section Report No. 7, 119 pp. BURDON, R.D. 2001. Pinus Radiata. In: Ecosystems of the World: Tree Crop Ecosystems. F.T. LAST (Editor), pp. 99-161. Elsevier.CSIRO, Canberra.)

Its cones are serotinous - closed until opened by the heat of a forest fire (surface fire); the abundant seeds - discharged to regenerate the burned forest. E.g. seedlings densities 1 year after fire - 100.000 up to 650.000/ha.

cold hardiness limit between -12.1°C and -6.7°C

principal host for the dwarf mistletoe Arceuthobium littorum

senescence - tree diameter reaches 100 cm(LITTLE, ELBERT L. Jr.. 1980. The Audubon Society field guide to North American trees. New York: Alfred A. Knopf.

BANNISTER, P. and G. NEUNER. 2001. Frost resistance and the distribution of conifers. P.3-22 in F.J. BIGRAS and S.J. COLOMBO (eds.), Conifer cold hardiness. Dordrecht: Kluwer Academic Publishers.

HAWKSWORTH, F.G. and D. WIENS. 1996. Dwarf mistletoes: Biology, pathology and systematics. Agriculture Handbook 709. Washington, DC: U.S.D.A. Forest Service. Available at World Wide Web: http://www.rmrs.nau.edu/publications/ah_709/index.html, accessed 15th October 2015. . WHITE et al., 1999. A nucleus breeding plan for radiata pine in Australia. Silvae Genetica 48: 122-133.)

ENVIRONMENTAL REQUIREMENTS AND ECOLOGY

CULTIVATION

natural stands - don´t appear to be single-aged!

breeding by seeds and cuttings (up to 15 years old specimen), 1 kg contains approximately 20 - 38.000 seeds

germination rate: 50 - 70%

germinate 20 days after sowing

seed storage: several years

seedlings: 4 - 8 - 12 (24) months old are planted

matrix: 1.5 x 1.5 - 3 x 3 m Mycorrhizae are necessary for seedling growth!

annual increment: 12 - 30 m³/ha, 2.5 m in height - S.W. England

rotation period: 20 - 50 years(WHITE et al., 1999. A nucleus breeding plan for radiata pine in Australia. Silvae Genetica 48: 122-133., Brink, M., 2008. Pinus radiata D.Don.

[Internet] Record from PROTA4U. Louppe, D., Oteng-Amoako, A.A. & Brink, M. (Editors). PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands. <http://www.prota4u.org/search.asp>. Accessed 19 October 2015.,

ÚRADNÍČEK, BROUMOVSKÝ and UNČOVSKÝ,1990. Lesní hospodářství v tropech a subtropech-Vybrané dřeviny, MZLU )

PEST AND DISEASES

Dothistroma needle blight (Mycosphaerella pini)

pitch canker –> the fungus Fusarium circinatum

Armillaria root rot

In South Africa the pine emperor moth (Imbrasia cytherea)

and the pine whoolly aphid (Pineus pini)PROTA (Plant Resources of Tropical Africa/Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands.(http://www.prota4u.org/protav8.asp?en=1&p=Pinus+radiata+D.Don)

GENETIC RESOURCES AND BREEDING

Chromosome number: 2n = 24

Abundant genetic variation –> highly successful breeding programmes.

Provenance testing and breeding –> South Africa and other major producing countries

Early breeding –> growth rate, tree form and disease resistance, now –> wood properties

Molecular biology –> genetic transformation of embryogenic tissue – biolistic and Agrobacterium-mediated systems, and stable transformed plants regeneration

PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands. http://www.prota4u.org/protav8.asp?en=1&p=Pinus+radiata+D.Don

Fig. 11 Diversity in cone size among the five native populations of Monterey pine (AXELROD, 1980)

Each cone represents the average size for that population. Key: 1 Cedros Island; 2 Guadalupe Island; 3 Monterey; 4 Año Nuevo; 5 Cambria

ROGERS, D. L. 2002.. In situ genetic conservation of Monterey pine (Pinus radiata D. Don): Information and recommendations. Report No. 26. University of California Division of Agriculture and Natural Resources, Genetic Resources Conservation Program, Davis CA USA. ISBN 0-9725195-0-5. Available on the World Wide Web: <http://ucanr.edu/repository/fileaccess.cfm?article=54987&p=DMIDEA&CFID=135918909&CFTOKEN=28218669>

Fig. 12 Summary of phenotypic characteristics of native populations of Monterey pine in field trials in New Zealand (BURDON 1992)

Symbols: + denotes superiority; – denotes inferiority; 0 denotes average; • denotes no data were located.

†Key: a denotes a large body of solid experimental evidence (many sites); b denotes good experimental evidence but from limited number of sites/pot trials; c denotes slender evidence; and two letters denote intermediate weights of evidence.

ROGERS, D. L. 2002.. In situ genetic conservation of Monterey pine (Pinus radiata D. Don): Information and recommendations. Report No. 26. University of California Division of Agriculture and Natural Resources, Genetic Resources Conservation Program, Davis CA USA. ISBN 0-9725195-0-5. Available on the World Wide Web: <http://ucanr.edu/repository/fileaccess.cfm?article=54987&p=DMIDEA&CFID=135918909&CFTOKEN=28218669>

Fig. 13 Allozyme diversity for the native populations of Monterey pine from three studies

Key: number of trees sampled per population (N), mean number of allele per locus (A), percent polymorphic† loci (P), and expected heterozygosity (He).

†With the exception of data from Plessas and Strauss, the criterion of polymorphism is 99%, meaning a locus must have a second allele with at least a frequency of 1% for that locus to be considered polymorphic. For the Plessas and Strauss data, the criterion is 95%, thus these data are an underestimate relative to the other data in the table.

ROGERS, D. L. 2002.. In situ genetic conservation of Monterey pine (Pinus radiata D. Don): Information and recommendations. Report No. 26. University of California Division of Agriculture and Natural Resources, Genetic Resources Conservation Program, Davis CA USA. ISBN 0-9725195-0-5. Available on the World Wide Web: <http://ucanr.edu/repository/fileaccess.cfm?article=54987&p=DMIDEA&CFID=135918909&CFTOKEN=28218669>

Fig. 14 Guadalupe Island pine seedlings in plantation

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G59100.html)

Fig. 15 Guadalupe Island pine plantation

(http://www.arkive.org/guadalupe-island-pine/pinus-radiata/image-G59628.html)

MAIN USESmedium hard wood and solid, less durable

sapwood: soft, creamy white, in case of young specimen holds 80 - 85% of volume

heartwood: pinkish brown, medium durable

wood density: (330–)380–610 kg/m³ at 12% moisture content

Construction wood, furniture, packing cases, poles, posts, shuttering, particle board boxes/crates, plywood, veneers, paper and fuelwood

Brink, M., 2008. Pinus radiata D.Don. [Internet] Record from PROTA4U. Louppe, D., Oteng-Amoako, A.A. & Brink, M. (Editors). PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands. <http://www.prota4u.org/search.asp>. Accessed 19 October 2015.

ÚRADNÍČEK, BROUMOVSKÝ and UNČOVSKÝ,1990. Lesní hospodářství v tropech a subtropech-Vybrané dřeviny, MZLU http://www.wood-database.com/lumber-identification/softwoods/radiata-pine/)

Fig. 16 Guadalupe pine wood

(http://www.prota4u.org/plantphotos/Pinus%20radiata%208.jpg)

Fig. 17 Guadalupe pine wood (sanded)

Fig. 18 Guadalupe pine wood (sealed)

(http://www.wood-database.com/wp-content/uploads/radiata-pine.jpg )

(http://www.wood-database.com/wp-content/uploads/radiata-pine-sealed.jpg)

MAIN USES

flooring, interior trim, toys, turnery, matches, railway sleepers, hardboard and wood-wool

oleoresin - distilled to obtain turpentine and rosin

turpentine - pine oil, terpene resins, flavours and fragrance

rosin - paper, inks, emulsifiers, synthetic resins, soap and glue

(https://books.google.cz/books?id=-nw-mZQ0kcEC&pg=PA445&lpg=PA445&dq=pinus+radiata,+subtropics&source=bl&ots=NPiikTfgUE&sig=lk-MFOXAq-VOWdm3oNWq3fndtEc&hl=cs&sa=X&ved=0CGQQ6AEwCWoVChMIp-63-erOyAIVhcByCh0DKw77#v=onepage&q=pinus%20radiata%2C%20subtropics&f=false)

RELATED SPECIES

all the species from genus Pinus (114), almost all are indigenous in the northern hemisphere

OTHER COMMENTSUse of radiata pine:

in windbreaks and as a shelter tree;

widely spaced trees with pasture beneath or as belts with pasture between them;

in woodlots, including stock havens

soil conservation

ornamental tree

may become invasive(http://www.fao.org/docrep/018/i3274e/i3274e11.pdf

https://books.google.cz/books?id=-nw-mZQ0kcEC&pg=PA445&lpg=PA445&dq=pinus+radiata,+subtropics&source=bl&ots=NPiikTfgUE&sig=lk-MFOXAq-VOWdm3oNWq3fndtEc&hl=cs&sa=X&ved=0CGQQ6AEwCWoVChMIp-63-erOyAIVhcByCh0DKw77#v=onepage&q=pinus%20radiata

%2C%20subtropics&f=false)

RESEARCH ARTICLES

Genome-wide gene expression dynamics of the fungal pathogen Dothistroma septosporum throughout its infection cycle of the gymnosperm host Pinus radiata (http://onlinelibrary.wiley.com.infozdroje.czu.cz/doi/10.1111/mpp.12273/full)

Effects of stand density and seedlot on three wood properties of young radiata pine grown at a dry-land site in New Zealand (http://nzjforestryscience.springeropen.com/articles/10.1186/s40490-015-0035-x)

Quantification of realised genetic gain in radiata pine and its incorporation into growth and yield modelling systems (http://www.nrcresearchpress.com/doi/10.1139/cjfr-2015-0191#.VttCTGDhCUn)

Metabolites and hormones are involved in the intraspecific variability of drought hardening in radiata pine (http://www.sciencedirect.com.infozdroje.czu.cz/science/article/pii/S0176161715002138)

Pattern of genotype by environment interaction for radiata pine in southern Australia (http://link.springer.com.infozdroje.czu.cz/article/10.1007/s13595-014-0437-6/fulltext.html)

Pattern of genotype by environment interaction for radiata pine in southern Australia (IVKOVÍC et al., 2014) – Introduction

Current radiata pine breeding and deployment in Australia is based largely on the plantation inventory zones rather than on biological patterns of genotype by environment interaction (G×E), and consequently cannot deliver optimal genetic gains across the whole plantation estate.

This study examined patterns of G×E to facilitate deployment of genetic stock to particular environments.

Pattern of genotype by environment interaction for radiata pine in southern Australia (IVKOVÍC et al., 2014) – Methods

20 genetically well-connected trials across southern Australia –> estimates of genetic correlations between performances at different trial sites.

Extended factor analyses (XFA) –> to estimate G×E variance and produce a matrix of site-site genetic correlations.

The patterns among these correlations – examined by heat map and hierarchical clustering.

Pattern of genotype by environment interaction for radiata pine in southern Australia (IVKOVÍC et al., 2014) – Results

The XFA captured a large proportion of both additive and non-additive G×E.

Significant G×E for diameter growth – expected between Tasmania and Mainland, and within Tasmania itself.

The study also confirmed presence of G×E between Murray Valley region in New South Wales and the rest of southern Australia.

Pattern of genotype by environment interaction for radiata pine in southern Australia (IVKOVÍC et al., 2014) – Conclusion

1. Based on previous studies and this study, significant G×E for diameter growth can be expected between Tasmanian and Mainland sites, and within Tasmania itself.

2. There were indications that the sites in Murray Valley region in NSW and Otway region in Victoria may exhibit G×E interaction with other regions; however, this is based only on individual trials.

3. Heritability increased significantly for within-region selection and regionalisation seems to be justified.

4. The G×E interaction at transcontinental scale can be correlated to the climate variables, primarily to rainfall and temperature. However, the drivers may also be related to smaller scale environmental variation (i.e. soil and terrain variation).

5. The results presented here can be used as evidence in favour of reconsidering the current breeding and deployment zones. However, further work, using other pairs of genetically connected trials, will give more robust results on which to base G×E regionalisation.

Metabolites and hormones are involved in the intraspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Introduction

Studies of metabolic and physiological bases of plant tolerance and hardening against drought –> essential to improve genetic breeding programs

Preliminary results: short drought period (4 weeks) –> different osmotic response in each breed

Acclimation in P. radiata – conditioned by the genotype + regulated by changes in physiological process and phytohormone concentration

Hardening – one of the most useful process for increasing plant drought tolerance and improve plantation success

Metabolites and hormones are involved in the intraspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Plant material

O1—P. radiata var. radiata × P. radiata var binata (Amberley, New Zealand)

O2—P. radiata var. radiata (Basque coastline, Spain)

O3—P. radiata var. radiata (Billapoola, Australia)

O4—P. radiata var. radiata × Pinus attenuate (Amberley, New Zealand) – tolerance marker

O5—P. radiata var. radiata × P. radiata var. cedrosensis (Amberley, New Zealand)

O6—P. radiata var. radiata (Kaingaroa, New Zealand)

Metabolites and hormones are involved in the intraspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Methods

Growth conditions – cold stratification, placing them into a cold chamber at 4◦C in dark for 3 weeks

–> seeds were placed in sterilized water for 2 days –same conditions to induce germination

–> seeds were sown in pots of 17 cm Ø filled with peat:perlite (7:3, v/v). Plants – greenhouse under controlled conditions (T = 23 ± 1◦C and RH = 70 ± 5%) for two years

Metabolites and hormones are involved in the intraspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Methods

Experimental design

Biometric, growth and water balance parameters

Water potential

Metabolite quantification – Free amino acids and polyamines and Hormone quantification

Statistical analysis – data compared by the parametric tests (2 and 3 way univariate analysis of variance-ANOVA) followed by Tukey’s test – open source R software 2.15.1. Normality – Shapiro’s test. No parametric data – Kruskal Wallis’ test

Metabolites and hormones are involved in the interspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Results: Biometric and physiological parameters

Different absolute growth – O1 and O5 the greatest total aerial height

O1 – the largest collar diameter

O5 – the highest values of relative height growth ratio(RdGR)

Water balance – no difference

Metabolites and hormones are involved in the interspecific variability of drought hardening in radiata pine (DIEGO et al., 2015) – Results: Metabolite content

Similar profile in plant hormones –> between breeds as hardening response, with significant increases in ABA, IAA, ZR, SA and JA and decreases in Z.

Exceptions:

O2 treated plants –> no change in their ABA and ZR content compared to controls

and O1 and O6 –> no significant differences in ZR and SA content, respectively.

Thanks for attention!