attack of the clones challenging new episode in our course ! clonal forestry dag lindgren
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ATTACK OF THE CLONES
Challenging new episode in our course !Clonal Forestry
Dag Lindgren
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Clones in Nature
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Clones dominated the early flora
0102030405060708090
100Number of families
% of flora
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Some large and old plant clones (after Cook, 1985). Species Diameter (m) Age (years) Gaylussacia brachycerium 1980 13 000 Holcus mollis 880 1000+ Populus tremuloides 510 10 000+ Pteridium aquilinum 489 1400 Lycopodium complanatum
250 850
Festuca rubra 220 1000+ Convallaria majalis 83 670+ Calamagrostis epigeios 50 400+ Picea mariana 14 300+ Festuca ovina 8 1000+ Larrea tridentata 8 11 000+
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Some natural clonal species in nature (Widén et al, 1994)
Species Genotypes per population (range)
Genotypes found in >75 % of populations
Alnus incana 5 100 Agrostis stolonifera 7,5 (1-15) 0 Antennaria rosea 3,5 (1-11) 0 Betula glandulosa 3 (1-5) 0 Erigeron annuus 14 (13-15) 59 Filipendula rubra 5,5 (1-15) 0 Glechoma hederacea 34,8 (15-98) 0 Oenothera biennis 1,5 (1-5) 0 Oenothera laciniata 6,5 (1-16) 1 Populus tremuloides 31,7 (30-40) 0 Pteridium aquilinum 24,8 0 Rubia peregrina 39,3 2 Rubus saxatalis 9 (8-10) 0 Solidago altissima 10,3 (3-20) 0 Taraxacum hollandicum 1,4 (1-4) 25 Taraxacum tortilobum 3,3 (1-7) 13
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Natural cloning in conifers is rather rareIn Scandinavian forest ecosystems clones
are most characteristic of the understory herbaceous vegetation, where they can be dominant
3-15 genotypes in a population represents a typical natural situation
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Clones in culture
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Examples of Agricultural Monoclonal Successes
‘Bartlett’ or ‘Williams’ peardiscovered 1770, 75% worldwide crop
‘Red Delicious’ appleorigin 1870, 50% worldwide crop
and many many more….
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From the history of clonal forestry
The first known use of vegetative propagation for forestry purposes was with Sugi (Cryptomeria japonica D. Don, a conifer) in Japan, where it is natural, in the 15th century.
Vegetative propagation of poplars (Populus sp.) has a long history in Europe. Organised clonal forestry started in the beginning of the 20th century
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Examples of clonal forestry
Species Area Magnitude
Eucalypts South America, South Africa, Portugal…
Around 15 million ha (half of plantations)
Radiata pine New Zealand 60 % of plantations
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Eucalyptus clones in Brazil
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Eucalyptus monoclonal block mosaic in Brazil
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Some reasons for a forester to use clonal forestry
To produce a more uniform product;To improve the forest by using a
genetically better planting stock;To get customer-tailored improved
material.
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A uniform product…
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Improved clones have advantages over improved seedlings!
Little time lag between selections in the breeding and practical forestry;
Short time span to deploy the gain achieved by breeding (get bulk of improved copies);
Flexible for variations in plant consumption;Eliminate problems with variations in seed
production;Eliminate problems in reproductive output or
reproductive phenology of trees in seed production unit;
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Possible advantages with clonal forestry
Eliminate selfing; Eliminate undesired unimproved pollen; Relatedness among selections little problem; The market for a bred material can be small Tailored varieties (no costumer too small...) Choose the level of diversity desired (often
uniformity) Mass-propagation of expensive but good seeds Possible to use non-additive variance (like
hybrid effects and dominance) Exploit the additive genetic variation better
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Possible advantages with clonal forestry
Choose clones which do not waist resources on sex
Create "physiologically programmed” types
Different cloning methods offer different options
Higher genetic gain (see above)Combine characteristics which seldom
are combined Test in different environments and choose for
wider adaptation (ecovalence)
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Comparison clone mix and seed orchard
Parameters as current Norway spruce breeding in Sweden
0
20
40
60
80
100
120
140
0 1 2 3 4 5 6 7 8 9 10 11
Generation (=20 years)
breeding pop
10 clone seed orchard
10 clone clonal mixture
Gain (per cent)
Rosvall, Lindgren & Mullin 1998
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Other reasons
Cuttings may be morphologically different, e.g. now insecticide use on conifer plants will be more restricted (Pyrmetrin); Cutting plants have thicker bark and more sturdy stem base, and are less harmed, and may possible constitute a solution. This may become the main reason for use of cutting Norway spruce plants in Sweden 2003.
It may be easier to make cuttings than to get seeds
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Often increased costs
More momentsClones must be conserved while testedLegal demands often multiply costsFor many species - expensive propagation
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Restrictions for clonal forestry
Experience over long time and large areas is needed, but accumulates slowly over time
Commercial and legal problems considerable and larger than with seeds
Get into problems with opinions, symbol of exploitation of Nature, "clone" is a dirty word
Not so profitable in practice as it appears from theory
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Risks and uncertainties Physiological state matters Propagation technique may matter for result Storage technique. Storing genotypes while
testing costs money and change characteristics. Somaclonal variations State of ortet may matter Juvenile age selection may give secondary effects Ecologic consequences? (probably limited) Specific pests and diseases may be favoured by
some clones and that they are repeated in stands and even spread from them
What happens at mature age? Often lack of mature field trials.
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How many clones?
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Genetic diversity in a stand is likely to favour production
A single genotype demands the same things at the same time, thus inefficient site use!
In a mix another genotype may take over the ecological space left by a failed genotype.
A disease or pest is expected to spread faster in a uniform crop.
Single clones have rather high G*E interaction thus may perform inexpectly bad under some conditions
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Clone number
Genetic and commercial
Gains
PlantationFailure
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Too much diversity in plantations?! Most crop- and many forest managers do not
like diversity Uniform trees means better economy and
simpler forestry even if biological production is lost
The genetic superiority of superior clones is much larger than the foreseen expected loss by uniformity.
The demand for “high diversity” in intensively managed forests may be very expensive in lost future gain
Uniform crops are easier to handle legally and commercially
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Better science possible
ReplicationsReproducibility
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A tool for a more effective breeding! Used for seed production Gains faster realized Clonal test means testing the sum of genes
deployed, progeny testing often are confounded by paternal genes just contributes to noise
A seedling is genetically unique; clones can be optimally replicated.
Clonal test gives in practice much information about ability to transfer gene to progeny
More efficient use of the variation occurring after sexual propagation
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A tool for a more effective breeding!
The efficiency of clonal testing depends on costs
Collaborators instead of competitors (non-egoistic clones)
Test on many environments and choose for wider adaptation
Test in the field and cross in the archiveCombine wood in the field and
reproduction in archiveBiotechnical breeding - like transgenetic
trees - becomes more feasible
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0
2
4
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16
0 500 1000 1500 2000 2500
Test size (plants)
Bre
edin
g v
alu
e
Clonal selection
Phenotypic selection
Comparison (at the same dimensioning) of clonal or seedling based testing for the Swedish Norway spruce long term breeding program. Clonal testing adds around 30% to gain. (Rosvall 1999)
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Cutting…
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Green-house for cutting production
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Clones may be made from somatic embryogenes
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Acknowledgements
Darius Danusevicius