Advantages of clonal propagation

DAG LINDGREN
The Swedish University of Agricultural Sciences, Department of Forest Genetics and Plant
Physiology, SE 901 83 Ume, Sweden
Last edit 01-10-27. Draft for the proceedings from a meeting at Ronneby, Sweden, August
2001.
Received November 1, 2001

Summary
Clones in forestry may be interesting for many reasons; three main purposes may be
mentioned:
To produce a more uniform product;
To improve the forest by using a genetically better planting stock;
To offer customer-tailored improved material.
Clonal propagation dominated in nature a long time ago (when plants entered land), and is
still common. Clonal Forestry appears in a large scale and since a long time (Sugi in Japan,
Eucalypts in Brazil and Poplars in Europe are examples).
Clones can increase the efficiency of forest tree breeding, even future seedling forestry
benefits from efficient cloning procedures.
Keywords: breeding, tree improvement, production population, clonal forestry,

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Introduction
Clonal forestry is often seen as modern biotechnology. It is true that some techniques used for
creating clones are new. The technical capacity to produce clones is improving. The current
paper aims to review the possible advantages of clones in forestry and forest tree breeding and
to give information about the occurrence of clones in nature and forestry. Some specific
attention is given to focus on matters, which are relevant for cloning as a tool to get quality
birch. No effort is done to discuss possible disadvantages and problems in full, but this has
been done many times before (e.g. Sonesson et al. 2001, Lindgren et al. 1990, Ahuja and
Libby 1993, Park et al. 1998). Neither are the possibilities of other competing alternatives
to get some of the advantages (like using controlled crosses) fully explored.

Clones in Nature and Forestry

Clones are common in Nature, and they have been used as a tool of domestication for very
long time and are used in large-scale forest operations. It seems to be important to remember
that to put clonal forestry into perspective,
The earliest land plants lacked vascular tissue and had inefficient systems of sexual
reproduction and dispersal and clonal reproduction was the typical way of propagation in
nature. Clonal growth was advantageous enabling physical dominance of large areas through
horizontal growth.
Clonal propagation is still important in nature for many species. Many of the commonest
clones are herbaceous but the habit is also found in woody plants throughout the world. In
temperate broad-leaved woodlands, root suckering is a common form of clonal growth and
there are good examples in the genera Populus and Prunus. Coppice is a natural growth form
that has been exploited for centuries in European forests. Betula, Carpinus, Corylus, Quercus,
Salix and Tilia are all genera with the ability to self-coppice.
The use of vegetative propagation of trees as a tool in their domestication has a long history.
An early use was to propagate good clones of fruit trees and this has been done for thousands
of years, thus humanity has long experiences of cloning trees.
For specific cases clonal forestry is often regarded as a new technique, and biotech leads
associations to the public to something more radical than most cases of clonal forestry. To put
things into perspective it has to be remembered that many of the techniques used in modern
seedling forestry are also new, and the problems and uncertainties with that is often larger
than the uncertainties connected to clones.
Some cases where clonal forestry has been much used are described, viz. Sugi, Eucalypts and
poplars.
Sugi
The first known use of vegetative propagation for forestry purposes was in Japan with Sugi
(Cryptomeria japonica D. Don, a conifer) in the 15th century. Since then reforestation with
monoclones or clonal mixtures has been widely used in Japan (see Ohba 1993). Cryptomeria
japonica is the most widely cultivated tree species in Japan comprising nearly half of the ten
million ha of Japanese plantations. Almost half of the plants used are clones (praxis differ in
different part of Japan and with different owners).
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The extreme uniformity of the plantations has probably contributed to greater than average
damage from typhoons and heavy snow. Attacks from Sugi bark borer and bark midge have
increased in severity as the area of Sugi plantation has grown, but there is no evidence of
altered disease or insect virulence related to clonal forestry, despite careful monitoring.
Clonal forestry is most common on the south-western island of Kyushu where about 100
cultivars are in use. Many foresters use only 1-3 cultivars of local origin in their plantations so
that a single clone may cover as much as 2 hectares.
Allergy to Sugi pollen is a health problem in Japan as a whole, but is only a minor problem on
Kyushu with its large use of clones. Cloning by cuttings for several generations has reduced
the incidence of male flowers, so these plantations produce little pollen. Analogously one may
speculate in that a reversion of North European birch forestry to clonal forestry in a very
remote future may reduce allergic reactions with birch pollen.
Eucalypts
Clonal forestry is more common with broadleaf species than with conifers. Probably the most
successful example is the eucalypt plantations in countries in tropical and subtropical climates
like South America, South Africa and Portugal. The total area of eucalypt plantations today is
about 30 million hectares and about half of this area is planted with cloned material.
The eucalypt plantations in Brazil cover an area of approximately 3 million hectares. A
succession of eucalypt species has been grown in Brazil due primarily to disease problems.
Clonal forestry started in the late 1960:s and is today the dominant form of eucalypt forestry
in the country. Progeny from an Eucalyptus grandis female (male sterile) growing in an
arboretum were found to be natural hybrids which were resistant to pests and diseases. The
offspring of this tree are believed to be Eucalyptus grandis Europhylla hybrids. This
knowledge was rapidly adopted by the forest industry companies and they selected clones
within the open-pollinated offspring from this single tree, which means that a small number of
clones originating from the same family are widely grown over large areas in Brazil. This has
and is still changing with most companies having good breeding programs backing their
clonal programs. The clones are deployed in huge monoclonal blocks. This situation seems
favourable for a pathogen, but nothing has happened yet - despite the possible risks of a
disease in a monoculture. Today the companies have become more risk-conscious and are
developing breeding and clonal testing programs that include strategies for clonal diversity in
space and time. A typical program is now deploying about 20 clones each year and these
clones will be replaced with time as new clones are selected. The production landscape forms
a genetic mosaic. New genetic material has been imported to widen the genetic base for
breeding.

Poplars and willows in Europe

Vegetative propagation of poplars (Populus sp.) has a long history in Europe. Organised
clonal forestry started in the beginning of the 20th century. Monoclonal plantations of poplars
have become a common land use especially on river plains in southern European countries
like Italy, Spain and France. Some of the best poplar clones in use were selected as early as
during the first half of the 20th century. Clonal forestry with poplars is common in countries
with subtropical and temperate climates like Italy, Spain, France, Belgium, USA and Canada.

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Single clones of poplars have been propagated extensively. There have been disease problems
and the overuse of single clones and use of large monoclonal plantations has been questioned
(cf e.g. Stelzer and Goldfarb 1997 p444). Poplar has covered over 100000 ha in one country
and a single clone may comprise one third of this area. The focus has been to develop disease
resistance within individual clones, but these clones have proved to be highly susceptible to
new varieties of disease. The disease problem seems to be growing and has caused Germany
to more or less abandon poplars and other countries may follow. Nevertheless these incidents
have not provided poplar growers with sufficient incitement to focus on more diverse
alternatives. It is unlikely that monoclonal cultures and the wide use of some clones are the
only explanation to increased disease, but they act as contributing factors. Legal and
commercial reasons often favour the use of single clones (e.g. easier to use breeders right
UPOV - for single clones than varieties, commercial demands to know that varieties are up to
specifications). Even willows and short rotation forestry in Sweden are threatened by disease.
Resistance is broken down within a single rotation (it is decades in between replanting even
when harvesting is done at intervals of a few years). Two evaluations of the Swedish willow
breeding program have rather strongly recommended an increased number of selections to be
made (more than a single selection released per year). Nevertheless, the growers
organisations still do not encourage clone mixtures.
Hybrid aspen in Finland
A Finnish forest industry company has recently started a clonal forestry program with hybrid
aspen (Populus tremula crossed with Populus tremuloides) for plantations in Finland and
Estonia. So far a few hundred hectares have been planted with micro propagated selected
clones. A recent Finnish doctor thesis has appeared on the subject of hybrid aspen (Yu 2001).
This raises the question if something similar could be done with birch. It may be noted,
however, that Yu (2001) suggested that micro propagation costs are high and that the
technique needs development.
Overoptimistic statistics
It is a clear tendency that statistics reported about the use of vegetative propagation, at least
there it is regarded as modern hitech biotech, gives the impression of more rapid progress than
actually occurs. E.g. Talbert et al. (1993, p. 148-149) reported, based on a survey, that four
Swedish operators produces 7.3 millions rooted Norway spruce cuttings annually. Lindgren et
al. (1990, p. 8) estimated the annual production to 5 millions and believed in a raise to 10
millions. However, looking backwards (Sonesson et al. 2001, p. 15), less than 20 million were
planted in total till 2000 and now only some hundred thousands are planted annually.

Crop uniformity
As genetic diversity and possible uniformity is an evident and possible important aspect of
clonal forestry. For evaluation of possible ecological impact diversity seems central. I will
thus start to discuss disadvantages with a low genetic diversity in a stand. In other respects
this paper focuses on the advantages of clonal forestry.
Disadvantages of a uniform crop
There are reasons to believe that the biological production can be higher in a genetically
diverse crop than in a uniform:
A single genotype demands the same things at the same time, and thus utilises the site
worse than a mixture of genotypes.
In a mix, another genotype may take over the ecological space left by a failed

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genotype.
A disease spreads faster in a uniform crop.

These statements are supported by the average of a very large number of agricultural mixing
experiments and a few experiments and experiences with forestry. On average, mixes
performed some percent better and were less susceptible and more stable than the average of
its components, but usually the production of the mix was not better than its best component
grown pure. Even if mixes seem more productive on an average, there are many examples and
experiments, when no advantage was found and some experiments where diversity actually
seems negative for production. I conclude that from a production point of view an advantage
with some degree of diversity is expected, but that does not imply that production is higher
the higher the diversity is. But the superiority in biological production is not expected to be
large and it is not certain for individual cases, but just a rough generalisation. This general
statement ought to apply to birch also, thus biological production is assumed to be lower and
more uncertain in a uniform crop, but the effect is probably small and practically unimportant
(a few percent of biological production).
Advantages of a uniform crop
There must be considerable advantages in uniform monoclonal blocks as they are used
frequently. Although the advantages with diversity are repeated and reiterated and very
politically correct, it is seldom applied in intensive agriculture or husbandry. The main reason
is that maximizing biological production and maximizing economical production is not the
same thing. It makes it easier to manage the crop if it behaves uniformly. For a forester it may
not matter very much if individual birches drop their leaves on different days, but a farmer
wants the crop to become ripe the same day, as the whole crop is harvested the same day.
Diversity is a usually a problem for the customer, who wants a uniform raw material for
further processing. For administrative processes, like describing the crop, it is an advantage if
it is uniform and does not change from specification over time and space. These problems are
evident for many types of agricultural crops, and therefore these crops are usually uniform
even when biological production would be higher with a diverse crop.
The dream of many foresters is to find the best genotype and use only that. This has been
successfully done for other domesticated species. The uniformity itself in a clonal plantation
has a considerable advantage from a management point of view. It is easier in the nursery to
raise plants of a single genotype. It is easier to manage and harvest a uniform forest. It has
advantages to handle a single type of material. A customer or processor of a product knows
more about a lot if it is a specified clone than if just the species is known. A batch of uniform
logs is worth more and easier to handle than the same amount of very variable logs.
Even if interest is only in quantitative production and that is lower in a uniform crop, still the
higher genetic performance of the best clones compared to the best seedlings available usually
much more than compensates for the production loss by a uniform crop. Thus uniform clonal
crops produce more than diverse seedling crops.
My opinion is that genetic diversity often is overemphasised in comparison with the higher
genetic gain achievable at the cost of a reduction in genetic diversity for future forest
production plantations in many countries (like Sweden) both for propagation with seedlings
and cuttings. To get genetic diversity many clones are used in seed orchards and clonal
mixtures and there constraints of relatedness of clones may be seen as desirable. This was not

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a disadvantage at an early stage of forest tree improvement, as it does not influence gain much
if many phenotypically selected plus trees are placed in seed orchards, and in the early stages
of cutting propagation there were physical constraints on the number of ramets which can be
produced by each clone, so it is necessary to have many clones. But now this constraint often
means selection is not intensive and the best genotypes are not exploited to the extent, which
would be possible.
However, clonal forestry is unlikely to deal with a single clone only. That does not seem a
likely scenario for birch. Many clones will be tested and test results as well as experiences
will accumulate of the best clones, so the opinion of what is the best clone will change over
time. It is also dependent on the locality, the desires of the user and the availability. Even if
uniform plantations are planted, there will usually be a mosaic of clones on the landscape
level. From the risk management point of view a clonal deployment philosophy offers
advantages. If a clone is sensitive to a pest or pathogen the problem can be identified and
managed. If silvicultural means to deal with a problem are insufficient, the stand can be
salvage harvested and replanted. The clone which turns out inferior can quickly be taken out
of production. It is an advantage for an operator which uses monoclone blocks to use several
clones in the same time that it will become more evident what the strong and weak point of
the clones are. A disadvantage with mosaics is that the uniform lots of wood from a stand are
often too small to be practical as a specific variety at the end user, and actually the end user
may get trouble with larger between lot differences if clonal forestry is used. Alternatively
the clones can be grown in intimate mixtures, in that they most of the possible advantages of
the diversity of seedlings may be gained by a mix of rather few clones.
When we are talking quality birch - and not quantity birch - the reasons for monoclonal
plantings become stronger. When high quality of the product is concerned, it often becomes
more important to have uniform quality also, while in a low quality product - as firewood the quantity is usually relatively more important and a uniform and predictable product is less
important. Attention on quality instead of quantity also generally coincides with a higher level
of intensity, and at a higher level of intensity, clonal forestry often becomes more interesting.
Thus considering the aim the project arranging this symposium has in mind; uniform clonal
plantings becomes relatively more important than for forestry in general.

Improved clones can be preferable to seedlings!

There are a number of arguments for forestry with clones. It takes less time to multiply a
genotype as a clone than to build a seed production unit with that clone as one of the parents
and wait till that works. The time lag is actually much longer than till the first seed
production, because a seed production unit is usually designed to last for many years. The risk
for getting the seed storage empty means that forestry often want to have five to ten years
foreseen seed need in storage. That is expensive, many seeds may never be used and the seeds
actually used are less improved than possible.
A seed production unit is designed to meet a certain need of plants with certain characteristics
many years ahead. If too many seeds are produced it means that the investment was
unnecessary large. If too few seeds are produced it may mean that foresters must use
genetically inferior unimproved seeds. If the propagation is done fast by vegetative
propagation by e.g. micro propagated clones or somatic embryogenesis, it takes only a year to

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multiply a desirable genotype in the desired number of copies. Thus it is fairly easy to adapt
to the latest news from breeders and customers.
There are large annual variations in seed production both in the forest and in seed orchards. In
a seed orchard the reproductive output is variable among genotypes and the reproductive
phenology may make clones incompatible, so the actual genetic set up of the progeny is a bit
uncertain. These uncertainties can be avoided by clonal forestry. Many of our forest trees like birch - can both spread pollen and produce seeds. That means that selfing can occur and
the occurrence of selfing in a plant material will decrease the forest production. With the use
of clonal forestry selfing can be completely eliminated.
In a wind pollinated seed orchard wild uncontrolled and unimproved pollen may contribute to
pollination, clonal forestry is a way to eliminate this problem. However, for birch, seed
orchards are small enough to be managed in plastic green houses, where the inflow of
unimproved pollen is low, so for birch it is not fair to see this pollen contamination problem
as an argument against open pollinated seed orchards.
In a seed orchard it is important to avoid related clones, as relatives get inbred progeny when
they mate, which means that forest production drops because of inbreeding depression.
Mating between relatives may be a worse problem than selfing in future seed orchards, as
self-zygotes usually die; self-seedlings are often culled before planting; or early out-competed
in forest, while milder inbreeding is more directly transferred to production loss. If clones are
related in a clonal forest, that does not cause inbreeding depression, and thus there is little
harm if related clones occur in a stand (actually this is very natural, trees in a natural stands
are often related). The clones in the breeding population will be increasingly more related as
breeding goes on, it will be easier to handle the consequences of this for the production forest
with clonal forestry.
A seed orchard is a big investment intended to meet the seed need for a long period for a
specific species used in a specified area, thus a considerable local market is needed to justify
the investment. Conifer seed orchards are usually designed for a need of many millions plants
per year over some decades. Multiplication of a clone can be seen as a smaller investment
during a shorter period, thus the local market is allowed to be much smaller with clonal
forestry, e.g. micro propagation for curly birch in Finland serves a market of 200 000
seedlings a year. A seed orchard has a stiff structure, its composition can be only slightly
modified once it is established and it is not easy to justify several orchards heading for
different characteristics for the same area. Clonal propagation is more flexible; an individual
customer can choose the clones, which fits the particular needs for the customer and occasion.
Sometimes a combination of rather rare characters is desired, a clone can be found which
combines several desirable characters in a much more efficient way than dealing with seed
orchards.
It is often stated that clonal forestry reduces genetic diversity. This is a generalisation, which
need not always be true. Actually, clonal forestry provides a tool to choose genetic diversity
at will, while when a seedling material is used, the genetic diversity is essentially left to
chance. Probably those using intensively managed birch plantation for producing high quality
timber with special characteristics would choose the option to reduce or eliminate genetic
diversity on the stand level, because they found it economically sound to do that. It seems
likely this is how the tool will be used. But clones in a clonal mix can be chosen so that the
genetic diversity is larger than in a seedling material. Clones can be mixed with seedlings,

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that may be a good strategy not only for diversity, but economically in cases where cloned
plants are much better, but also much more expensive. If the clones succeed well, their share
at the final harvest will be larger than at establishment. Over time and space different clones
will be used even in a system with monoclone cultures (like Eucalypts in Brazil). In such a
system the genetic diversity among stands is likely to be much than in nature. Sometimes
clones have been used for natural conservation in situation when good seeds could not be
obtained. Clonal forestry may strengthen the trend to produce the needed wood at a limited
acreage of intensively managed production forests, allowing larger areas to be set aside for
other purposes. Thus the question how clones may affect diversity has many aspects and it is
an oversimplification to state that it must be severely reduced.
A common situation is that some good seeds exists or can be obtained, but quantitatively
insufficient for the plant need. If such a situation occurs unexpected and unforeseen,
vegetative propagation may be the solution. A more common situation is that a few superior
seeds can be produced by controlled crosses, but too few to meet the plant demand. Clonal
propagation is often a way to make forestry with controlled crosses practical feasible. Clonal
propagation may be seen as a way to amplify the impact of a few good parents or a tested
cross by mating the parents and when multiply the seeds.
The genetic value of a genotype depends on the breeding value of its parents, but also how
well the genes of the parents fit together. This may be called dominance. The later reason for
genetic differences cannot be exploited in an ordinary seed orchard, but it can be exploited
with clonal forestry.
Hybrids sometimes result in good seeds. It may be difficult to make many of those seeds by
controlled crosses, but hybrid seeds (or trees) can be multiplied by vegetative propagation.
This is used in the Eucalypts and Aspen programs mentioned above. Personally I am not that
convinced that hybrid superiority is so common as it is often stated. When hybrids are better
than the parents it may be because the parents were not well adapted for the conditions where
they were tested or that the hybrid combines characters of the parents for the environment and
purpose of the forester in a better way than tested alternatives. Hybrid maize is a best seller,
but the reason is more that farmers are unable to produce hybrid seeds them self, but has to
buy them, and when it is good economy of a seed company to invest in hybrids, so they can
market the seeds.
To get a genetically superior forest, it is desirable to use the genes of the very best clones. In a
seed orchard it has to be rather many parents to avoid selfing and problems with lack of
overlap in flowering time. This means a drop in breeding value. For clonal forestry the very
best clone can be used. This is true even if in the complete absence of dominance. As
dominance is seldom the major cause of genetic variation for the most important characters in
forest tree breeding, this more effective exploitation of breeding value is usually a more
important reason for clonal forestry than the exploitation of dominance variation, at least as
long as we do not talk about hybrids of different species or races. Quantitatively the low
importance for the superiority of cloning of a limited amount of dominance variation is
demonstrated by e.g. Lstiburek (2000).
We have also special features for clonal selection. We can intentionally select clones, which
do not waist resources on sexual activities but spend resources on stem wood production.
Such clones are desirable for forestry. In a seed orchard it is not desirable with clones, which
do not produce seed and pollen, so such clones would not be chosen for a seed orchard, and if

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they were, they would be unable to transmit their genes to the seed crop.
Cloning methods have different characteristics. They are often very depending on the state of
the material which is cloned, cuttings usually do not work well on physiologically old
material, for somatic embryogenesis other tissue than immature seeds are often a difficult
starting material. With micro propagation it is usually possible to multiply old trees or clones,
and this is often an advantage.
Cloning and the cloning method can have effects on the physiological status of the tree. E.g.
conifer grafts have not thick bark at the bottom of the trunk. The protocol for making clones
can have effects on the plants. These effects sometimes look desirable and can be exploited.
For example Norway spruce cuttings seems to suffer a lower mortality from pine weevil
damage than seedlings (Mattson and Thorsn 1992). Clones usually origin from older and
more mature tissues than seedlings and this affects the characters of the plants and may have
effects also on the mature trees. I guess these differences to seedlings are not dangerous and
nursery raised seedlings can also be claimed to be unnatural, and they my be different because
of different growing regimes. I suggest that we can intentionally interfere with the characters
of the plants by the production system and that clonal forestry gives additional options to use
this in a positive way.
Many of the factors I have mentioned above will contribute to that clonal forestry gives a
genetically more improved forest than comparable seedlings. An example of the development
of gain over time for the clonal option versus the seed orchard option is shown in Figure 1.
This is based on a simulation of the Swedish Norway spruce (Picea abies K.) long term
breeding program with inputs chosen to correspond to the real program as close as possible.
Clonal forestry and seed orchards are different ways of harvesting gain from the breeding
population and clonal forestry means that the genetic improvement reaches a certain level
Swedish Norway spruce breeding

Gain (per cent)

140

120

100

Clonal forestry
80

60

Breeding pop
40

Seed orchard

20

0
0

10

Generation (=20 years)

three decades before the same level can be achieved by seed orchards.

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11

Figure 1. A comparison of the genetic value of three populations as a function of time based on a
simulation of the Swedish Norway spruce breeding program. Genetic progress is primarily
created in the breeding population. The production population is created by creaming the best
available material from the breeding population either by clonal forestry or seed orchards. The
gain unit can roughly be interpreted as percent gain. The details of the figure depend on exact
specifications, but it gives an idea of magnitudes based on best possible estimates. Clonal
forestry gives a higher gain than seed orchards; the difference at a given time is in the magnitude
of 15%. The same gain is achievable from seed orchards; it just takes a few decades longer to get
it. Modified from Rosvall, Lindgren and Mullin 1998, Figure 1.

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More effective breeding with clones!

In the section above it was pointed out that clonal forestry might be a faster way to better
forests than seed orchards. Good genotypes are multiplied as grafts in most current forest seed
orchards, thus clones are useful also for producing genetically improved seedlings. But there
is another - still more powerful - way of using the clonal tool to improve the genetic quality of
forests. Clone testing has the potential to make the long-term breeding considerable more
efficient. Even if clones are too expensive for direct use in forestry, their use in testing of
genetic materials may help forestry to get better seeds in the future. A combination of testing
clones for forestry and for long-term breeding has synergistic effects, thus such a program is
more efficient than a program focusing only on one of these two aspects.
Higher organisms are diploid, thus have two set of homologous chromosomes and carry two
packages of genes. The parents only transfer one package (a haploid gamete) to the progeny,
and the composition of that gamete is different for each progeny. Thus, if offspring is used to
test parents, only half of the genes in the offspring are relevant for the parent, and these genes
will be different for each offspring individual. In contrast, in clonal testing all genes in the
genotype are tested and no genes generate random noise in the statistical sense.
The performance of a clone depends on its father, its mother and how the genes of the mother
and father match together, thus dominance effects. However, dominance variation does not
seem large enough for most characters of practical interest to be important (e.g. Rosvall 1999,
Lstiburek 2000). Thus the performance of a clone in a clone test says much about what will be
inherited to its progeny.
The accuracy by which the breeding value of a clone is determined increases with the number
of ramets. The uncertainty caused by the environment decreases then environment is repeated.
Thus clonal testing is a way of increasing the accuracy. One can see it as a way of increasing
the heritability to decrease the environmental variance by letting the environment be the
average of a number of individual environments, one for each ramet. The gain obtained by a
genetic selection depends on the selection intensity and the accuracy of the test. Under the
constraint of a fixed plant number, a lower number of genotypes can be tested if genotypes
are replicated by cloning, and thus the selection intensity will decrease It thus becomes an
optimization problem. However, if the individual heritability is not very high and the resource
budget is not extremely tight, the optimum is to clone in at least some copies.
A seedling material is genetically diverse; the best genotypes grown pure are much better than
the average of a seedling material. The performance of a seedling is mainly depending on its
environment and not so much of its genes. If it is cloned, the average over the clone is more
depending on the genes. Testing by clones can be seen as a way to eliminate the statistical
noise. The heritability of clonal values will be much higher than the heritability of seedling
values.
Each seedling is genetically unique; a scientific principle is to use replications when testing.
Thus a seedling is a rather unsatisfactory material for genetic testing and conclusions based on
seedlings will be dependent on model assumptions, which are uncertain and affected by
statistical noise. Clone testing is less depending on model assumptions and a more direct
measure and thus offers theoretical advantages!

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In a series of investigations (e.g. Rosvall 1999; Danusevicius and Lindgren 2002; assuming
conditions similar to the Swedish Norway spruce program), it has turned out favorable to test
the breeding value of the candidates for the breeding population by clonal testing instead of
phenotypic testing or progeny testing, if that is at all possible.

16
14
Clonal selection

Breeding value

12
10
8

Phenotypic selection

6
4
2
0
0

500

1000
1500
Test size (plants)

2000

2500

Figure 2. Comparison of clonal or seedling based testing for the Swedish Norway spruce long
term breeding program (the X-axes gives the resource level, the test size for the real program
is around 560 plants/parent). Clonal testing adds around 30% to the gain of the breeding
program, the advantage is larger the more resources that are available for each parent (Rosvall
1999).
The superiority of clonal testing over phenotypic testing is demonstrated in Figure 2 (from
Rosvall 1999 Figure 3). It is based on the same type of simulations of the Swedish Norway
spruce long term breeding program as the previous figure, although it varies the resources
available (thus the size of the tests). The resources available today correspond to 560 test
plants per parent. The more resources available, the more reason to use clone testing to get a
more accurate selection (compared to increasing the number of test genotypes for a more
intense selection).
Lstiburek (2000) and Danusevicius and Lindgren (2002) studied the conditions under which
clonal testing is a favourable alternative. The results show that clone test is superior to
phenotype test and also progeny test under a wide range of conditions. Thus even if the
genotypes or ramets are expensive; and over a wide range of available resources; and in the
presence of typical dominance variation; clonal testing can be recommended. In cases where
the individual heritability is close to 1, clonal testing is less favourable. When the phenotype
itself gives sufficient information about the genotype, the accuracy need not be improved by
cloning. Such cases are rare for growth characters, but may be more common for some quality
characters.

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Numerous studies have shown that environment can cause lasting changes in phenotype that
can be passed from one generation to the next, much as genes are transmitted (Schwaegerle,
McIintyre and Swingley 2000). Such affects may limit the usefulness of clonal testing for
breeding value, but similar objections could be made against progeny testing, and the effects
must anyway be larger than usually observed (e.g. Lindgren and Wei 1994) to really matter.
Natural genotypes are not designed for perfect cooperation; the Darwinian selection in the
struggle for life unavoidable gives some preference to strong competitors. A certain degree of
selfishness is an unavoidable consequence of natural selection. By clonal selection we ought
to be able to identify and utilise the best collaborators, who act in the interest of the stand and
not only the individual tree. Here is a case when Man may have the means to do better than
Nature and identify collaborative clones.
A forest material is used over varying conditions in time and space and the forester wants a
reasonable stable material over these variable conditions. Clones can be tested over time and
space. That make breeders able to select clones, which perform well everywhere and anytime.
Natural selection works just here and now. Again a case where Man may find more effective
ways than Nature has used.
Tree Breeders first select based on field tests and when mate the selections. Artificial crosses
are difficult to organise in the remote places where field tests are localised. But if clones are
tested, some copies could be placed where it is convenient to make artificial mating, like in
archives outside the institute.
Foresters do not want trees to waist resources on sex, but to do breeding it is necessary to get
reproduction to work. But clones, which have little reproductive output in the field, can have
copies at the research station, which are manipulated to make mating possible. Thus with
clones different individuals can be used for test and reproduction.
Genetically modified trees will play a role in practical forestry and tree breeding at some time
in the future. Such implementation is much easier, if clonal forestry is an established praxis.

Conclusion
If cloning is economically feasible and biologically practical, where are strong genetic reasons
for using it in practical forestry as well as long-term tree breeding.
Acknowledgements
Some of the work reported here (Rosvall 1999; Lstiburek 2000; Danusevicius and Lindgren
2002) as well as this paper itself has been supported by the EU project FAIR5 PL97 3823
Wood quality in Birch.

References
Ahuja, M.R. and W.J. Libby (Eds.).1993. Clonal Forestry I: Genetics and biotechnology,
Clonal Forestry II: Conservation and application. Springer Verlag. Berlin.
Danusevicius, D. and D. Lindgren. 2002. Comparison of Phenotypic, Clonal and Progeny
Supported Selection in Long-term Tree Breeding. Silvae Genetica in press.
Lindgren, D., Gustafsson, L., Hulthn, H. and K Lundkvist. 1990. Klonskogsbruk. SLU.

Dag Lindgren; November 2001 File: DagBirchProceedingsMs0202

Page13

Skogsvetenskapliga fakulteten. Rapport nr 9. Ume. (In Swedish)

Lindgren, D. and R.-P. Wei. 1994. Effects of maternal environment on mortality and growth
in young Pinus sylvestris field trials. Tree Physiology 14:323-327.
Lstiburek, M. 2000. Theoretical analyses of the possible benefit of vegetative propagation for
quality birch. Examensarbete i Biologi/Skogsgenetik. Printed at Department of Forest
Genetics and Plant Physiology. Swedish University of Agricultural Sciences.
Mattsson, S. and Thorsn. 1992. Regeneration and its inherent problems. In The
Rationalisation Conference 1992. G. Frumerie (ed). Redogrelse No 1. The Forest
Operations Institute of Sweden, Stockholm, pp 3548. (In Swedish with English
summary).
Ohba, K. 1993. Clonal Forestry with Sugi (Cryptomeria japonica). In Ahuja, M.R. and W.J.
Libby (Eds.). Clonal Forestry II: Conservation and application. Pp 66-90. Springer
Verlag. Berlin.
Park, Y.S., Bonga, J.M., and T.J. Mullin. 1998. Clonal forestry. Chapter 8 In Forest genetics
and tree breeding. Edited by A.K. Mandal and G.L. Gibson. CBS Publishers, New Delhi,
India. pp. 143-167.
Rosvall, O. 1999. Enhancing Gain from Long-Term Forest Tree Breeding while Conserving
Genetic Diversity. Acta Universitatis Agriculturae Sueciae. Silvestria 109 65pp+4
chapters.
Rosvall, O, D. Lindgren, and T.J. Mullin. 1998. Sustainability, robustness and efficiency of a
multi-generation breeding strategy based on within-family clonal selection. Silvae
Genetica, 47:307-321.
Sonesson, J., Bradshaw, D. Lindgren and P. Sthl. 2001. Ecological evaluation of clonal
forestry with cutting-propagated Norway spruce. SkogForsk Report 1: 59 pages.
Stelzer, H.E. and B. Goldfarb. 1997. Implementing clonal forestry in south eastern United
States. Can J For Res 27:442-446.
Schwaegerle, K.E., McIintyre H, and C. Swingley. 2000. Quantitative genetics and the
persistence of environmental effects in clonally propagated organisms. Evolution 54:452461.
Talbert CB, Ritchie GA and Gupta P. 1993 Conifer Vegetative Propagation: an Overview
from a Commercial Perspective. In Ahuja, M.R. and W.J. Libby (Eds.).1993. Clonal
Forestry I: Genetics and biotechnology, pp. 145-181.
Yu, Q. 2001. Selection and propagation of hybrid aspen clones for growth and fibre quality.
University of Helsinki. Department of Applied Biology. Publication 6. Doctoral thesis. 41
pp and 5 chapters.

Dag Lindgren; November 2001 File: DagBirchProceedingsMs0202

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