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Polyploidy – A tool in adapting trees to future climate changes?

A review of polyploidy in trees

Ræbild, Anders
Elsevier Scientific Publ. Co
2024-05-15

Ræbild , A , Anamthawat-Jónsson , K , Egertsdotter , U , Immanen , J , Jensen , A M ,

Koutouleas , A , Martens , H J , Nieminen , K , Olofsson , J K , Röper , A C , Salojärvi , J ,
Strömvik , M , Vatanparast , M & Vivian-Smith , A 2024 , ' Polyploidy – A tool in adapting
trees to future climate changes? A review of polyploidy in trees ' , Forest Ecology and
Management , vol. 560 , 121767 . https://doi.org/10.1016/j.foreco.2024.121767

http://hdl.handle.net/10138/576298
10.1016/j.foreco.2024.121767

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 Forest Ecology and Management 560 (2024) 121767

Contents lists available at ScienceDirect

Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Polyploidy – A tool in adapting trees to future climate changes? A review of

polyploidy in trees
Anders Ræbild a, *, 1, Kesara Anamthawat-Jo
!nsson b, Ulrika Egertsdotter c, Juha Immanen d, e,
Anna Monrad Jensen , Athina Koutouleas a, Helle Jakobe Martens a, Kaisa Nieminen d, e, Jill
f

Katharina Olofsson a, Anna-Catharina Ro”per g, Jarkko Saloja

”rvi e, Martina Stro
”mvik h,
i j, 2
Mohammad Vatanparast , Adam Vivian-Smith
a
University of Copenhagen, Department of Geosciences and Natural Resource Management, Rolighedsvej 23, Frederiksberg DK-1958, Denmark
b
Institute of Life and Environmental Sciences, University of Iceland, Sturlugata 7, Reykjavík IS-102, Iceland
c
Swedish University of Agricultural Sciences, Dept. of Forest Genetics and Plant Physiology, PO Box 1234, Umeå SE-901 83, Sweden
d
Production Systems, Natural Resources Institute Finland (Luke), Helsinki 00790, Finland
e
Organismal and Evolutionary Biology Research Program, Faculty of Biological and Environmental Sciences, Viikki Plant Science Center, University of Helsinki, Helsinki
00014, Finland
f
Department of Forestry and Wood Technology, Linnaeus University, V” axj”o 351 95, Sweden
g
Center for Bioresources, Danish Technological Institute, Taastrup 2630, Denmark
h
Department of Plant Science, McGill University, 21111 Lakeshore road, Sainte-Anne-de-Bellevue, QC H9X 3V9, Canada
i
Jodrell Laboratory, Royal Botanic Gardens Kew, Richmond, TW9 3AE, United Kingdom
j
Norwegian Institute of Bioeconomy Research (NIBIO), Division of Forestry and Forest Resources, P.O. Box 115, Ås N-1431, Norway

A R T I C L E I N F O S U M M A R Y

Keywords: Polyploidy, or genome doubling, has occurred repeatedly through plant evolution. While polyploid plants are
Adaptation used extensively in agriculture and horticulture, they have so far found limited use in forestry. Here we review
Ecophysiology the potentials of polyploid trees under climate change, and investigate if there is support for increased use. We
Fitness
find that polyploid trees like other plants have consistent increases in cell sizes compared to diploids, and that
Forestry
leaf-area based rates of photosynthesis tend to increase with increasing levels of ploidy. While no particular trend
Tree breeding
Whole genome duplication (WGD) could be discerned in terms of biomass between trees of different ploidy levels, physiology is affected by poly-
ploidization and several studies point towards a high potential for polyploid trees to adapt to drought stress. The
ploidy level of most tree species is unknown, and analysis of geographical patterns in frequencies of polyploid
trees are inconclusive. Artificial polyploid trees are often created by colchicine and in a few cases these have been
successfully applied in forestry, but the effects of induced polyploidization in many economically important tree
species remains untested. Polyploids would also be increasingly useful in tree breeding programs, to create
synthetic hybrids or sterile triploids that could control unwanted spreading of germplasm in nature. In
conclusion, this review suggests that polyploid trees may be superior under climate change in some cases, but
that the potential of polyploids is not yet fully known and should be evaluated on a case-to-case basis for different
tree species.

1. Introduction Hartmann et al. 2022), reforestation could have a significant impact by

regaining control of the terrestrial carbon cycle, and help counteract the
Forest ecosystems contribute significantly to the diversity and pro- effects of climate change. A simple but so far unexplored method to
duction of the Earth’s biomass (Bar-On et al., 2018). While there is enhance the robustness and productivity of many forest tree species
concern for the future health of forests due to climate change (e.g., could be polyploidy, or genome duplications. Since the 1930’s, natural

* Corresponding author.
E-mail address: [email protected] (A. Ræbild).
1
ORCID 0000-0002-8793-5663
2
ORCID 0000-0002-8074-7950

https://doi.org/10.1016/j.foreco.2024.121767
Received 20 December 2023; Received in revised form 9 February 2024; Accepted 12 February 2024
Available online 15 March 2024
0378-1127/© 2024 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

polyploids, induced mutations and polyploidization by chemicals and 2021). Trees, however, differ from non-woody plants in having sec-
irradiation have been used for general plant improvement (Darrow, ondary growth and extended xylem. This makes trees tall and wide,
1950; Sigurbjornsson and Lachance, 1987), shaping agriculture and giving them an edge in ecosystems when competing for light. In addi-
leading farmers into the ’Green revolution’ (Hedden, 2003; Jung and tion, trees have longer generation times and life spans than herbs and
Till, 2021). Although many examples of polyploidization in agriculture forbs, which may affect their potential for adaptation to rapid envi-
and horticulture have led to significant improvement in productivity ronmental change. Given the hypothesized evolutionary advantage of
and trait performance (Motosugi et al., 2002, Wu et al., 2012), its polyploids during previous environmental upheavals (Van de Peer et al.,
application in forestry has been limited. However, there are many po- 2017), polyploids may have an adaptive advantage in their response to
tential advantages of utilizing polyploids in forestry including increased the challenges of climate change (Lomax et al., 2014; Faizullah et al.,
resistance to abiotic stresses (e.g., Maherali et al., 2009; Tossi et al., 2021; Ebadi et al., 2023).
2022), superior performance under elevated CO2, and changed wood There is extensive research on the evolutionary role of poly-
properties (da Silva Souza et al., 2021). ploidization and whole genome duplications (WGD) in key terrestrial
Polyploids often have altered plant development compared to their plant radiation events, particularly for tracheophytes (Soltis et al., 2015;
diploid relatives. Key phenotypic differences, such as larger cell sizes for Clark and Donoghue, 2018). Even as this review was concluding, Bure#s
higher ploidy levels, tend to hold in ploidal series across plant orders et al., (2024) explored the relationships between bioclimatic environ-
including bryophyte, fern, gymnosperm and angiosperm lineages ment, genome size, polyploidy and growth form in angiosperms, and
(Henry et al., 2014; Scholes and Paige, 2015; Alix et al., 2017; Karlin and concluded that woody angiosperms were less frequently polyploid and
Smouse, 2017; Smarda
# et al., 2018; Wilson et al., 2021; Patel et al., that genome size regulation may operate differently. There is, however,

Box 1
Glossary/list of terms.

Accessory or cloud genes: Genes that are found in some, but not all, species or individuals in a taxon or taxonomic group. These genes are seen
as non-essential, however, they bring uniqueness and are often advantageous to the bearer (e.g. disease resistance genes, stress tolerance,
flowering time).
Allele-dosage/gene-dosage: The variable amount of gene product that is the result of multiple copies of an allele, or gene, on homologous and
homeologous chromosomes.
Allopolyploid: A polyploid formed from the merging of gametes from different species or highly diverged populations.
Aneuploidy: State of organisms and cells with an unbalanced set of chromosomes with respect to their parents. Aneuploid organisms may have
chromosomal instability (chromosome breakage causing gene loss) or have an abnormal number of chromosomes, such as one extra or one fewer
chromosomes than expected.
Apomixis: Is the process of clonal reproduction via asexual seeds, and is often found associated with natural polyploids and ploidal variation.
While apomixis often has complex evolutionary origins, and often low frequency amongst plants, the molecular basis for naturally occurring and
synthetic modes of apomixis is now well understood, and may provide significant benefits to agriculture or forestry due to the fixation of hybrid
vigor.
Autopolyploid: A polyploid formed by whole genome duplication (WGD) often caused by merging of unreduced gametes from individuals of
the same species.
Copy number variation: The variation in the number of genes for discrete loci caused by tandem duplication, or from the number of gene loci
occurring in polyploids.
Core genes: Set of genes that are shared by all individuals or species in a taxon or taxonomic group.
Diploid: Cells of an organism that have two complete sets of chromosomes, one set from each parent. Chromosomes are represented as ho-
mologous pairs in diploid organisms, and the diploid state is typical of somatic cells.
Genomics: The study of the structure, function, and evolution of the total genetic and/or epigenetic sequence information (the genome) of
organisms.
Haploid: Cells of an organism that have one complete set of chromosomes, typically gamete cells.
Heterosis: Non-additive inheritance of phenotypic traits as observed from hybridization of two different parental lines, species or genera.
Hybrid vigor is a highly sought after effect of heterosis.
Homeologous chromosomes: matching chromosomes in the genomes of different species. Previous to speciation, these were homologous
chromosomes.
Homologous chromosomes: the matching chromosomes in two sets of chromosomes in diploids, or in more than two sets of chromosomes in
autopolyploids.
Neo-functionalization: The development of a new function of a gene copy (co-option) following duplication.
Neopolyploid: A newly formed polyploid lineage. Neopolyploids can be further defined as neoallopolyploids and neoautopolyploids.
Polyploid: Cells of an organism that have more than two complete sets of chromosomes. Depending on the number of chromosome sets, they are
referred to as triploid (three sets), tetraploid (four sets), pentaploid (five sets) etc.
Sub-functionalization: Gene copy expression patterns associated with different or novel time or tissue types following gene duplication.
Sub-genome: The set of chromosomes derived from distinct progenitor species.

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A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

a continued lack of understanding of the specific role of polyploidy for genomes of allopolyploids have a conserved order (synteny) for the
forest tree species during evolution. This review therefore examines if majority of their core genes, each sub-genome can also contain a unique
the benefits of altered plant development that are often seen in polyploid set of genes (accessory or cloud genes) that may confer stress or disease
crop plants, also hold for polyploid forest trees. We summarize the resistance to newly formed polyploids (neo-polyploids) (Chen et al.,
contemporary research on natural and artificial polyploid forest trees to 2020). Neo-polyploids, in particular neo-allopolyploids, may thus have
investigate whether there is evidence for the superior performance of important advantages over their diploid progenitors. Hybrids and
polyploid trees under climate change conditions, and if polyploidy can neo-allopolyploids may also display heterosis and thus outperform their
be used as a tool in forestry and breeding programs to enhance the parental lineages. The genomic underpinnings of heterosis, however, are
resilience of our forests. not very well understood, but are suggested to be controlled by a com-
plex set of molecular processes including sub-genome dominance
2. Formation of polyploids and molecular effects of (Carlson et al., 2022; Carlson and Smart, 2022; Yu et al., 2021) and
polyploidization haplotypic divergence, changes in allele-dosage and novel regulatory
interactions affecting gene expression networks (Hu et al., 2022). Ex-
Polyploidization is an integral part of plant evolution, and most, if amples of the latter are found in Salix L. (Carlson et al., 2022; Carlson
not all, plant genomes have undergone polyploidization at some stage and Smart, 2022), Populus L. (Liqin et al., 2019), and Quercus L. (2 n →
(Van de Peer et al., 2017; Simonin and Roddy, 2018). While whole 36; Butorina, 1993; Dzialuk et al., 2007; Johnsson, 1946; Lefort et al.,
genome duplications within a species, autopolyploidization, can 1998; Lefort et al., 2000; Naujoks et al., 1995). The fast growth of these
happen, allopolyploids are thought to be more common. Allopolyploids trees could be derived from a combination of copy number variation and
are established from hybridization between divergent lineages followed heterosis.
by genome duplications (Box 1, Fig. 1; Otto and Whitton, 2000; Mallet,
2007). Early research on (allo)polyploids provoked a hypothesis on a 3. Changes in anatomy and morphology of polyploid plants
“Genomic shock” (McClintock, 1984) in the neo-(allo)polyploid species,
possibly driven by transposable elements, with extensive genome-wide There is a tendency across plant species that cell size correlates with
changes. In contrast, recent research has found a peaceful coexistence genome size (Cavalier-Smith, 2005; Hodgson et al., 2010; Robinson
of the sub-genomes, with the regulation of different processes divided et al., 2018), although this may be cell type dependent (Katagiri et al.,
between them (e.g., Bird et al., 2018; Duan et al., 2023). Over time 2016). Increased cell size affects a great number of key physiological
polyploid genomes tend to become re-diploidized (Ramsey & Schemske, processes in plants such as water holding capacity, and transport of
2002; Conant et al., 2014). Following the initial polyploidization event, gasses and solutes (Trojak-Goluch et al., 2021).
the novel lineage undergoes genome fragmentation (Zhu et al., 2019), Since polyploid plants tend to have larger stomata that are posi-
either with equal gene loss in both sub-genomes (Liang and Schnable, tioned further apart compared to their diploid relatives (e.g., Smarda
#
2018), or with one sub-genome losing genes at a more rapid rate while et al., 2023, Fig. 2), this trait can be used to identify polyploids. This is
the other becomes dominant, as in Gossypium (Chen et al., 2020; Con- also true for tree species across a large phylogenetic spectrum (Fig. 3a,
over and Wendel, 2022). This process can also result in novel genetic b). Increases in stomatal cell size may be accompanied by increases in
variation in the form of neofunctionalization or sub-functionalization, the size of other cell types, including epidermal cells and mesophyll
where multiple gene copies provide the opportunity for further parenchyma, resulting in thicker leaves in polyploids (Chen et al., 2021).
specialization (Liang and Schnable, 2018; Van de Peer et al., 2021). Past This means that both the amount of dry mass and the water mass per leaf
ancestral history of polyploidization may also have a persistent influ- area may increase in polyploids compared to diploids and has been re-
ence on gene expression and genomic dominance (Yoo et al., 2013; ported in several woody genera (Barcelo !-Anguiano et al., 2021; Li et al.,
Liang and Schnable, 2018). 2009; Romero-Aranda et al., 1997; Zhang et al., 2019a). Larger cell sizes
While the molecular and genomic effects of polyploidization are well in polyploids do not necessarily lead to a larger surface area of leaves, as
studied in particular crop plants, little data has so far been obtained for there may be a correspondingly lower number of cells in the leaves
tree species due to their long generation times and thus difficulties in (Robinson et al., 2018). Data from woody species are contrasting, with
multi-generational studies. The molecular effect of hybridization, and some studies showing that polyploids have shorter, but thicker leaves
allopolyploidization, depends on the genetic similarities between the than their parental diploids (Xue et al., 2017), some showing no increase
diploid progenitors, their divergence time and differences in their gene in leaf area (Diallo et al., 2016; Fernando et al., 2019), and some
regulation, with perhaps the most extensive effects (i.e. genomic shock) showing larger leaf areas in polyploids (Chen et al., 2021; Hao et al.,
resulting from more diverged progenitors. For example, while the sub- 2013; Wang et al., 2017b). The number of studies is too limited to relate
these differences to phylogeny or habitats of the species.
Sizes and numbers of organelles are also affected by the level of
ploidy. Larger stomata are associated with an approximate doubling of
chloroplast numbers in guard cells in tetraploids compared to diploids
(Beck et al., 2003; Tang et al., 2010; Shi et al., 2015; Abdolinejad and
Shekafandeh, 2022), and chloroplast sizes were approximately double in
tetraploid Mangifera indica L. compared to diploids of the same species
(Barcelo !-Anguiano et al., 2021).
There are relatively few studies that go beyond leaf anatomy, but
they seem to indicate that polyploids have larger cell sizes in other parts
of the plant. For example, tetraploid Citrus genotypes have been shown
to have thicker roots than diploids (Syvertsen et al., 2000; Ruiz et al.,
2016a). Polyploids had larger structures in the hydraulic systems such as
longer and wider vessel elements and sieve tube elements (Hao et al.,
2013; (Barcelo !-Anguiano et al., 2021; De Baerdemaeker et al., 2018)). In
Fig. 1. Natural formation of polyploids from diploids (2x) to triploids (3x), species of Betula L. from China vessel diameters were larger in polyploids
tetraploids (4x) and hexaploids (6x). The figure illustrates autopolyploids from compared to diploids, although intervessel pits had smaller dimensions
whole genome duplication, and allopolyploids via inter-specific hybridization in polyploids, resulting in a larger resistance to cavitation compared to
and back-crossing from e.g. two different species. Modified from Yirga (2021). diploids (Zhang et al., 2017).

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A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

Fig. 2. The sizes of stomata are positively correlated to genome size in Acacia senegal. a) Tetraploid, stomata length 22.2 ↑ 1.4 µm. b. Diploid, stomata length 16.8 ↑
1.2 µm. n→ 10. Bar→ 20 µm.

Fig. 3. Anatomical and physiological traits in tree species with varying ploidy levels. a) Stomatal density. b) Stomatal length. c) Operating stomatal conductance, Gs.
d) Net photosynthesis, Pn. Inserted P-values indicate tests of effects of ploidy level from a mixed model analysis with the ploidy level as a fixed factor and species/
variety as a random factor. Data and references are available from Table S1.

Floral morphology and anatomy of trees are also affected by ploidy 4. Physiology of polyploid trees
level, with polyploids showing overall larger morphologies than diploids
(Borges et al., 2012; Fernando et al., 2019; Oliveira et al., 2022). This 4.1. Photosynthesis in polyploid trees
includes pollen (Shi et al., 2015, Mendes-Rodrigues et al., 2019) and
polyads as well as stigma cups (Diallo et al., 2023). It is generally sug- It has been hypothesized that smaller genomes are advantageous
gested that such differences could act as a reproductive barrier for gene under low CO2 conditions because this leads to smaller leaf structures
flow between ploidy levels, thus contributing to reproductive isolation. and thus faster diffusion of CO2 to the sites of photosynthetic assimila-
Although the available evidence suggests larger cell sizes throughout tion (reviewed by Faizullah et al., 2021). This theory is corroborated by
all organs of the polyploid plants compared to diploids, there is insuf- reports of extensive reduction in genome size of angiosperms during the
ficient data to verify whether all cell types are affected to the same de- Cretaceous, which allowed for reductions in cell sizes, higher intracel-
gree by ploidy. As a larger cell size may influence physiological lular surface areas and more efficient CO2 diffusion into photosynthetic
performance and wood quality, there is a need for additional studies of mesophyll cells, in turn maximizing primary productivity (Simonin and
the anatomy of polyploid trees at several levels of the plant. Roddy, 2018; Th! eroux-Rancourt et al., 2021). Polyploidization in trees

4
A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

may, however, have been most relevant during the Devonian and part of examples of polyploid superiority (compared to diploids of the same
the Mesozoic, known for high humidity and atmospheric CO2 concen- species) include conditions of chilling stress, bacterial disease and
trations (Veselý et al., 2020). Under these conditions, larger (slower) salinity (Ruiz et al., 2016a,b; Luo et al., 2017; Zhao et al., 2017; Sivager
stomata would not have been detrimental to the overall CO2:H2O status et al., 2019, 2021; Lourkisti et al., 2020). Here we focus on climate
of the plant, as they could remain open for longer periods without change where drought has been identified as a major challenge.
risking excessive water loss (Veselý et al., 2020). However, with the Stress trials have shown evidence of drought resilience in polyploid
climatic shifts occurring during the Carboniferous to Permian periods trees (Diallo et al., 2016; Lourkisti et al., 2021a,b). The underlying
and from the mid-Cenozoic onwards, the cooler and drier environments biochemical and signaling mechanisms which cause greater drought
(characterized by low atmospheric CO2 concentrations) may have tolerance in polyploids are not well understood. However, it has been
incurred a selective pressure on stomatal downsizing to optimize shown that the important abscisic acid (ABA) signaling pathways are
intracellular CO2:H2O (Veselý et al., 2020; Wang et al., 2021). Yet, the affected (Allario et al., 2013; Rao et al., 2020). In addition, osmotic
only study to test this hypothesis in trees found that increases in net adjustment and other water-saving physiological traits (in polyploid
photosynthesis under elevated CO2 were similar in tetraploid and trees) could explain the apparent drought tolerance compared to diploid
diploid genotypes of the genus Citrus L., indicating no superiority of individuals (as is the case for Betula - Li et al., 1996; Malus - Zhang et al.
tetraploids in this context (Syvertsen et al., 2000). 2015; de Baerdemaeker et al., 2018; Citrus - (Lourkisti et al., 2021a,b;
Although very little is known about how diffusion inside the leaf is Lourkisti et al., 2022; and Mangifera indica - Perera-Castro et al., 2023).
affected by changes in ploidy levels, the ploidy-induced, leaf anatomical Significantly higher concentrations of soluble sugars, proline, and
changes may modify the diffusion pathway for CO2 and water vapor. glycine betaine were found in tetraploid genotypes compared with
These leaf traits can potentially alter conductance and possibly even the diploid genotypes of young Ficus carica L. (Abdolinejad and Shek-
CO2:H2O ratio. For example, larger leaves in polyploids may have afandeh, 2022). Accumulation of these compounds no doubt plays a role
greater boundary layer resistances than smaller leaves of lower cyto- in plant cell-water balance.
types (Nobel, 1999). Stomatal conductance (gs) in polyploids is reduced Changed xylem anatomy in polyploids also appears to lead to
by a generally lower number of stomata per leaf area, but on the other changed hydraulic properties (Barcel! o-Anguiano et al., 2021; De Baer-
hand could be increased by their larger pore sizes compared to diploids demaeker et al., 2018; Hao et al., 2013). Root hydraulic conductivity of
(Faizullah et al., 2021, Smarda
# et al., 2023). Our analysis of differences tetraploids was found to be smaller compared to diploids in both Citrus
in gs among ploidy levels for a range of tree species shows no general ssp. and Salix viminalis L. (Dudits et al., 2016; Ruiz et al., 2016a,b), but in
tendencies (Fig. 3c). Thus, ploidy-induced anatomical changes to the other Citrus genotypes, the hydraulic conductivity was not affected by
stomatal apparatus may affect conductance and its regulation capacity. the level of ploidy (Syvertsen et al., 2000). In a modeling approach
Although not directly testing for the effect of ploidy, intracellular where polyploids were assumed to have larger vessel and sieve tube
diffusion of gasses and water have been shown to be slower in large sizes, polyploids showed increased hydraulic conductance compared
mesophyll cells and enlarged chloroplasts compared to smaller and more with diploids (Barcelo !-Anguiano et al., 2021). Additionally, De Baer-
densely shaped anatomies (Xiong et al., 2017; Th! eroux-Rancourt et al., demaeker et al. (2018) found higher relative water content and lower
2021). In Acacia senegal (L.) Willd. and Populus tremuloides Michx., stress response in autotetraploid under in-vitro drought conditions linked
carbon isotopic composition suggested that water use efficiency is to gene expression levels for aquaporin controlling genes (MdPIP1;1 and
greater in polyploids than in diploids, as would be expected when the MdTIP1;1). However, the hydraulic conductance of the shrub Atriplex
diffusive pathway for CO2 is longer (Greer et al., 2018; Diatta et al., canescens (Pursh) Nutt. decreased with increasing ploidy despite larger
2022). vessel sizes (Hao et al., 2013). This suggests that hydraulic conductance
The larger dimensions of leaves, cells and organelles in polyploids cannot be evaluated based on vessel anatomy alone. Importantly, Hao
are mostly associated with increases in rates of net photosynthesis (e.g., et al. (2013) found that plants with higher ploidy had smaller loss of
Xue et al., 2017; García-García et al., 2020; Chen et al., 2021). A limi- hydraulic conductivity under tension, suggesting that polyploids would
tation of several studies is that they are based on one or two genotypes of be more resistant to severe drought.
each cytotype, restricting conclusions to the specific genotypes involved. Upon water deficit conditions, vascular plants initiate systemic
However, our analysis across different species and genotypes indicates intercellular communication involving the phytohormones ABA and
that photosynthesis (when expressed on a leaf area basis) is higher with cytokinins, which eventually accumulate in the leaves and cause sto-
increasing levels of ploidy (Fig. 3d). Still, Populus triploids had faster matal closure (Verslues, 2016; Takahashi et al., 2020). This reduces
rates of photosynthesis and growth compared to diploids and tetra- additional water shortages, which could occur via leaf transpiration. The
ploids, suggesting that there is not always a direct link between genome ABA in tetraploid Prunus salicina Lindl. was found to accumulate later
size and physiology (Zhao et al., 2015; Zhang et al., 2019a). than in the diploid leaves under water-stress (Pustovoitova et al., 1996),
There is a scarcity of studies examining the scaling effects, which attributed to the larger cells and organelles of the polyploid, thus
come with larger genome sizes and how this interacts with plant CO2 imposing a biophysical constraint on the signaling of ABA. However, the
uptake and water loss (Wang et al., 2021). The ‘large genome constraint final ABA content and cytokinin activity in the tetraploid leaves were
hypothesis’ was tested across phylogenies in the context of photosyn- found to exceed that in the diploid leaves. Although the transpiration
thetic performance, demonstrating that photosynthetic rates (per leaf rate decreased more rapidly in tetraploids under drought conditions it
weight) were lower with increased genome size in gymnosperms, but recovered more quickly after subsequent rehydration episodes
not in angiosperms (Knight et al.,2005; Beaulieu et al., 2008). We are compared to diploid plants, suggesting an acclimation response to the
not aware of studies investigating this dynamic in polyploid trees. Since drought stress exhibited solely by the tetraploids after repeated drought
thick leaves may have high rates of photosynthesis (when expressed on a exposure (Pustovoitova et al., 1996).
per-area basis) and low rates (when expressed per dry weight), a pros- Collectively, these controlled and semi-controlled drought experi-
pect lies in the possibility that methods used to estimate photosynthesis ments in growth chambers and greenhouses suggest the superior per-
may lead to opposite conclusions regarding photosynthetic capacity in formance of polyploid trees under drought. Unfortunately, this has
polyploid trees. rarely been tested in field trials. Studies under controlled conditions and
in the field of diploid and triploid Populus tremuloides have come to
4.2. Physiological variation of polyploid trees under drought opposite conclusions, pointing to an overall lower resilience to drought
in triploids under field conditions (Greer et al., 2018; Blonder et al.,
There is a limited number of studies on the stress responses of 2021; Eisenring et al., 2023). Despite greater instantaneous water use
polyploid trees, mainly focusing on horticultural species. However, efficiency, triploid P. tremuloides in the field showed lower resilience to

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A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

climate-induced stress and had higher mortality than diploids (Blonder growth and cell enlargement, and thus their altered functions and
et al., 2021). Studies such as these include extraneous variables but are expression in the autotetraploid could explain the altered phenotype
better indicative of the eco-physiology of polyploid trees under natural (Mu et al., 2012b). A similar increase in biomass was reported for au-
environments compared to laboratory assessments. totetraploids of the fast-growing deciduous species of Paulownia elongata
S.Y.Hu (Zhi-qing, 2006). Several genes were shown to co-regulate
biomass production and are thus presented as ideal targets in breeding
4.3. Regulation of growth in polyploid trees programs (Cao et al., 2020). Amongst others, genes involved in
improved chloroplast formation, and thus improved photoassimilation,
The “gigantism” exhibited by the autotetraploids can be associated were put forward to be most likely responsible for the increased biomass
with altered signal transduction of indoleacetate (IAA) and upregulation in the tetraploid (Cao et al., 2020).
of six genes involved in the biosynthesis and signal transduction of There are, however, contrasting results between studies and species
ethylene (Mu et al., 2012b). The autotetraploid Betula platyphylla concerning the regulation of growth. Some researchers suggest that
Sukaczev has a unique phenotype compared to diploids of the same plant growth is reduced with increasing ploidy level due to increased
species, with a more dense stature characterized by increases in numbers of chromosomes, which may lead to slow mitotic division, and
breast-height diameter, volume, leaf and catkin sizes. Also, certain lower overall growth (Ramsey and Schemske, 2002). In particular,
triploid Betula lineages were found to be superior in base diameter and meiotic and mitotic rates have been hypothesized to be slower in poly-
height to base diameter ratio when compared to the diploid lineages (Mu ploids (compared to diploids) due to more time needed for the
et al., 2012a). Both IAA and ethylene have key roles in regulating plant

Fig. 4. a) Heat map presenting the distribution of tree species per country based on the GlobalTreeSearch database (Beech et al., 2017); b) Polyploid tree species
distribution heat map generated based on the dataset from Rice et al. (2019). Note: areas with low frequencies may be under-represented in this database.

6
A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

replication of the genome prior to cell division (Tate et al., 2005). diversity; for example the Dipterocarpaceae, a tropical tree family that is
Another possible explanation for slow cell division in polyploids is the foundation species in many forest ecotypes in South East Asia, and is
spatial constraints (Corneillie et al., 2019), particularly concerning the the major source of high quality hard wood in timber trade in the region,
organization of chromosomes during metaphase alignment and spindle consists of around 700 species (Appanah and Turnbull, 1998).
attachment, which are more likely to be perturbed or delayed due to the Polyploidy can be detected by direct or indirect methods. Indirect
high number of chromosomes in the nucleus of polyploid species methods rely on the analyses of morphological traits such as larger
(Comai, 2005). Slow ontogeny has also been reported in tetraploid po- stomata in lower density and/or a higher number of chloroplasts per
tato (Solanum tuberosum L.) (Tamayo-Ordo !n
$ ez et al., 2016). However, guard cell (see above). In contrast, direct methods measure the nuclear
these delays in development could be mitigated by faster and early genome size by flow cytometry, which measures DNA content per cell,
germination by larger seeds, which is a common characteristic of poly- and/or chromosome counting, both of which are more accurate than
ploid plants (Tate et al., 2005; Selvi & Vivona, 2022). Growth in poly- indirect methods. In particular, chromosome numbers obtained visually
ploid trees may also be limited by the increasing costs of larger cell size from microscopic images provide further information, such as karyo-
structures, for example pollen tube growth (Williams & Oliveira, 2020). typic variation and genome composition in polyploids, and offer deeper
This feature was observed in both Betula and Handroanthus species. insights into chromosome evolution especially when using molecular
There may also be relevant interactions between genetic background cytogenetic techniques.
and ploidy level which may be overlooked, if single genotypes are Estimating the number and distribution of polyploid tree species
examined. Genotype-ploidy dependent interactions have been demon- globally is challenging due to the lack of chromosome counts for most
strated in Zea mays L., where the morphological response to ploidy tree species. Globally, chromosome records are available for less than
change was genotype specific and shown to vary along inbred lines 20% of angiosperms and are biased toward specific clades (Rice et al.,
(Riddle et al., 2006). 2015). Unfortunately, only 10% of all tree species have available chro-
Given the lack of consistency between studies, there is still a need to mosome numbers in the Chromosome Counts Database (data retrieved 1
examine the role which genome size plays in the regulation of growth in April 2023). Advanced techniques like estimating genome size with flow
trees. Until this is understood, the potential use of polyploid trees in cytometry are used, but actual genome size data is available for only a
forestry must be studied individually for key species and genotypes. fraction of plant species (Pellicer et al., 2018). To increase our knowl-
edge, prioritizing affordable chromosome counting methods is crucial,
5. Natural occurrence of polyploidy in trees especially in tropical and subtropical regions where tree species di-
versity is concentrated.
Trees are vital for global plant biodiversity, representing about There are, however, some genera, including birch (Betula), that have
60,000 species and nearly 20% of seed plants (Beech et al., 2017). The been extensively studied. Betula comprises 40–60 tree/shrub species
highest diversity of tree species is found in the tropical regions of South distributed from the sub-tropical and temperate to the arctic regions of
America, Asia, and Africa (Fig. 4a). Estimates suggest that 30–70% of the Northern Hemisphere (eFlora, 2023, Furlow, 2023). Hybridization
angiosperms and approximately 10% of gymnosperms are polyploids among Betula species is frequent (Ashburner and McAllister, 2013;
(Wood et al., 2009; Rastogi and Ohri, 2020; Ohri, 2021). In general, Johnsson, 1945; Walters, 1964) and hybrids are often found in the
polyploid frequency tends to increase with distance from the equator, overlapping area of natural distributions. In contrast to oak (Quercus),
influenced by climate, particularly temperature (Ramsey and Schemske, beech (Fagus L.) and chestnut (Castanea Mill.) that are ploidy-stable at
1998; Rice et al., 2019; Bure#s et al., 2024). In contrast, polyploid tree the genus level (mostly diploid 2 n→2x→24; Chokchaichamnankit et al.,
species appear to be more common in tropical regions (Fig. 4b). This 2008; Rice et al., 2019), Betula is an example of a ploidy-variable genus
heatmap of distribution is likely biased, however, as it is based only on a (Fig. S2, Table S2). Among Betula species, the ploidy levels range from
few thousand ploidy-known tree species worldwide. The exact number diploid (2x) to dodecaploids (12x), with the tetraploid number
of polyploid tree species and the extent of natural occurrence of poly- 2 n→4x→56 being the most common (Rice et al., 2015; Wang et al.,
ploid tree species remains unknown. 2016; eFlora, 2023). Overall, there are more polyploid (56%) than
In gymnosperms, polyploids are rare and mainly found in Ephedrales diploid (44%) species, and in terms of abundance and geographic range,
and Cupressales, with high incidences in Ephedra L. (76%) and Juniperus polyploid species are more common in Eurasia than in North America
L. (22%) (Ohri, 2021). In addition, there are stable polyploid cultivated (42 vs. 15 species; Table S2). Interestingly, the proportion of polyploid
forms of Ginkgo biloba L. (Smarda
# et al., 2018), and the hexaploid Betula species does not increase at higher latitudes as is generally
Sequoia sempervirens (D.Don) Endl. is one of the most valuable timber observed among other flowering plants. Several arctic and alpine species
species in the forest industry in Northern America and New Zealand are diploid, such as the dwarf birches B. nana L. and B. glandulosa Michx.
(Meason et al., 2016). In angiosperms, frequencies of polyploids appear (de Groot et al., 1997) and Alaskan tree birch B. neoalaskana Sarg.
to be variable among taxonomic clades (Fig. S1, S2). At the family level, (Furlow, 2023), whereas a few sub-tropical Asiatic species are polyploid,
Fabaceae, Asteraceae, Malvaceae and Rosaceae have the highest such as B. alnoides Buch.-Ham. ex D.Don and B. cylindrostachya Wall.
numbers of polyploid woody plant genera (Fig. S1a). Forest tree genera (both tetraploids with 2 n→4x→56, eFlora, 2023). In addition, the ratio
with a relatively high proportion of polyploids, such as Alnus Mill., Tilia of diploid to polyploid Betula species is the same in both mountain
L., Sorbus L., and Betula, are those having their natural distribution range forests at high altitudes and in the lowland areas (Table S2). Thus,
in the northern temperate region (Fig. S1b,c). The genus Sorbus (rowan) diploids and polyploids of Betula are symmetrically distributed. In the
comprises some 150 species mostly in the northern temperate regions. case of Alder (Alnus species), most polyploid species are distributed in
These include mostly out-crossing diploids (2 n→2x→34), apomictic the East Asia and Mediterranean region, whereas European and Amer-
triploids (2 n→3x→51) and tetraploid (2 n→4x→68) species ican species are predominantly diploid (Vít et al., 2017). Recently
(Hajrudinovi!c et al., 2015; Kaplan et al., 2019; McAllister, 2005). Most discovered tetraploid populations of A. glutinosa (L.) Gaertn. in Morocco
of these polyploid species have deficient fertility, having originated from (Lepais et al., 2013) and the newly described tetraploid species
interspecific hybridization, and then followed by WGD in the case of A. lusitanica Vít, Douda & Mand! ak and A. rohlenae Vít, Douda & Mand! ak
tetraploidy. Therefore apomixis, which is defined as the clonal repro- in Southern Europe and North Africa (Vít et al., 2017) open possibilities
duction via asexual seed (Underwood and Mercier, 2022; see Box 1), to test the performance and potential use of these taxa in forestry.
provides a mechanistic advantage that allows the continued survival of
triploid individuals within this genus. Since there is still relatively little 6. Producing artificial polyploid trees for forestry
genomic research on tropical tree species, the proportion of polyploids is
largely unknown. This is also hampered by the immense species Different methods to obtain chromosome set doubling have been

7
A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

applied to in vitro cultures as a fast-track method of obtaining artificial or even sometimes self-incompatibility are used to manage pollination
polyploids of commercially interesting tree species like Eucalyptus L′H!er. of female acceptor lines with the tetraploid pollen sources. Hybrid
(Silva et al., 2019), and also to restore fertility in newly synthesized production in this manner usually avoids, or provides a way to tolerate
alloploids (Sattler et al., 2016). Artificial polyploids in trees have been endosperm imbalance which can arise from interploidy crosses during
most frequently induced by the antimitotic agent colchicine, which hybrid seed development (Gehring and Satyaki, 2017).
mechanistically disrupts the cell cycle (Touchell et al., 2020). Different Production systems that include triploidy in forestry may require
tree parts can be treated with colchicine to induce chromosome significant time for their development and assessment. Likewise, their
doubling, the first step in polyploidization. However, most treatments appropriateness may need to be coordinated for forest biodiversity or
target different types of in vitro cultures of shoot tissues, leaf material the perceived risk from an invasive fertile polyploid forest tree species
and established meristematic tissues. Mature leaf blades of Populus x which may have a significant growth advantage (e.g. hybrid eucalypt
hopeiensis Hu & H.F.Chow were treated with colchicine and the induced species). Even so, polyploidization in forest breeding programs may
tetraploid plants mostly came from around vein tissues (Wu et al., 2020). provide other useful opportunities. Synthetic polyploid systems have
Furthermore, octoploid plants were derived from the tetraploid plants, been used to overcome breeding barriers, and perform wide hybridiza-
and the phenotypic consequences of gene dosage effects were evaluated tions (Manzur et al., 2015; Ruiz et al., 2020), or regain fertility (Riese-
in the ploidy series of 2x, 4x and 8x plants (Wu et al., 2022). Allohex- berg, 2001). While fertility is often unaffected in induced
aploids were also generated in another effort to investigate the perfor- autopolyploids, since endosperm balance is unchanged during seed
mance of economic traits in Populus with higher ploidy levels (Liu et al., development, synthesized allopolyploid hybrids can have reduced
2018). In Eucalyptus polybractea R.T.Baker, nodal shoot cultures were fertility. Often the restoration of fertility occurs through further
successfully treated with colchicine to induce formation of thicker doubling of allopolyploids, and can be attributed to a number of specific
leaves with larger oil glands (Fernando et al., 2019). Seeds were steril- mechanisms (Rieseberg, 2001). These include modifying the ploidal
ized and treated with colchicine to induce chromosome doubling in balance between subgenomes, balancing the divergence between the
Acacia dealbata Link and A. mangium Willd. (Blakesley et al., 2002). The parental subgenomes, or eliminating of specific genomic sequences, and
tetraploid plants were subsequently used for breeding to generate sterile improving the stability of the hybrid genome. In other cases,
triploid trees. Colchicine treatment of seeds was also successful for post-zygotic atrophy can occur at the F1 generation (Manzur et al.,
inducing tetraploids in Platanus x acerifolia (Aiton) Willd. (Liu et al., 2015), and fertile individuals are sometimes obtained by embryo rescue.
2007). In horticultural tree species, like citrus and mandarin, shoot Even considering those barriers, there are many examples where syn-
apical meristems (SAM) were treated ex situ, and after shoot induction thetic polyploids have acted as a bridge between species of different
grafted as scions onto trifoliate rootstocks (Aleza et al., 2009; Ollitrault ploidy levels, allowing new genetic diversity, via introgressive hybrid-
et al., 2020). Such a method avoids the requirement for the introduction ization, to be tapped into plant and tree breeding programs.
of material into in vitro culture that may pose considerable challenges for Changed ploidy levels can sometimes be associated with clonal
recalcitrant tree species or long-lived woody perennials. reproduction via asexual seeds or apomixis (eg. Sorbus; Box 1). This is
More powerful classes of antimitotic agents, such as the herbicides often observed at low frequency in tree species, and often unlikely to
dinitroanilines (trifluralin and oryzalin) and phosphoric amides (ami- have significant impact on population structure, with the exception of
prophos-methyl and butamiphos; Planchais et al., 2000) are not widely Mangifera, Citrus and Sorbus for example, where obligate modes of
used to induce chromosome set doubling in trees. Oryzalin was however apomixis can and do occur (Hamston et al., 2018; Wang et al., 2017a;
used with some success to induce polyploidy in Eucalyptus dunnii Yadav et al., 2023). The molecular basis for apomixis is now well un-
Maiden, but the survival rate was low (Castillo et al., 2020). derstood (Underwood and Mercier, 2022), and engineering of apomixis
into different crops has been recently achieved (Vernet et al., 2022;
7. Potential for polyploidy in forest tree breeding Underwood et al., 2022). Similar to triploid breeding programs,
apomixis offers the potential for the fixation of hybrid vigor, and a
The induction of polyploidy in crop species often creates opportu- method to evaluate morphological plasticity and niche adaptability.
nities and changes in breeding systems. Triploid breeding programs can Furthermore if asexual seed reproduction were ever to be introduced
be particularly beneficial, however there have been only few instances into key forestry species, individuals with polyploid genomes may be
of implementation and evaluation in key forestry species, e.g., Acacia better suited for the expression of both parthenogenesis and modifier
(Nghiem et al., 2018), Populus (Kang and Wei, 2022), Eucalyptus uro- loci (Hojsgaard, 2018). Apomixis on the other hand, will not provide a
phylla S.T.Blake (Yang et al., 2018) and Betula (Mu et al.,2012a). suitable method of germplasm containment for an elite breed of forestry
Naturally occurring triploids have also been identified and selected for trees should that be necessary.
clonal propagation due to their potential for enhanced traits and gain
(Quercus, Dzialuk et al., 2007; Prunus avium L., Serres-Giardi et al., 2010; 8. Uses of polyploids in forestry
Eucalyptus grandis x urophylla, Longui et al., 2021). The low fertility of
triploid oaks, likely due to disrupted meiosis and pollen production with The many examples of accelerated vegetative growth in polyploids
unbalanced chromosome complements, means their direct value for (Beyaz et al., 2013, Chen et al., 2021) have led to an interest to test the
downstream breeding is low (Butorina, 1993). Overall, however, it potential of both natural and artificial polyploid trees for forestry pur-
would appear that the advantages of triploid breeding in forestry pro- poses. While examples of polyploid trees are numerous in horticulture,
grams have not yet been fully realized. In contrast, horticultural triploid the number of natural polyploid tree species that are currently in use in
breeding programs are advanced and provide significant utility. For a forestry is relatively limited. Still, the list of currently ploidy-known
large variety of crops elite diploid and tetraploid parental lines are bred trees and shrubs in Table S3 contains commercially used species,
to produce hybrid seed, almost all of which are triploid. These progenies including many shrub species but also important forestry species such as
offer many benefits including plants with a fixed heterotic growth Acer pseudoplatanus L., Fraxinus chinensis Roxb., and several tropical
advantage and a uniform phenotypic distribution, along with sterility timber genera such as Shorea Roxb. ex C.F.Gaertn. and Cordia L. Trees
and the absence of seeds (Sattler et al., 2016). Having triploid progeny that naturally display multiple levels of ploidy are predominantly found
has the added advantage of containing the germplasm and avoids the amongst angiosperm trees and provide valuable material for testing
release of elite breeding lines with stacked traits including disease effects of ploidy on field performance, like Acacia senegal (Diallo et al.,
resistance into the natural environment. Horticultural seed production is 2016) and Populus tremuloides (Arno and Hammerly, 2020, Blonder
often managed using crosses onto male diploid sterile acceptor lines et al., 2021). Betula (Table S2) contains several valuable polyploid
(Cheng et al., 2022). In this case, nuclear or male cytoplasmic sterility, species, including B. chinensis Maxim., which is one of the most valuable

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A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

timber trees in northern China (eFlora, 2023), and the North American than the diploids (Fernando et al., 2019).
paper birch (B. papyrifera Marshall, comprising tetra-, penta- and Since gymnosperm trees, in particular species within the genera
hexaploids), one of the most sought after wood tree species for building Pinus L., Picea Mill. and Larix Mill., includes several commercially
and other purposes (Furlow, 2023). In northern Europe, the tetraploid important species, efforts have been made to produce superior traits
B. pubescens Ehrh. is a common species that has previously been bred for through the generation of colchicine-induced polyploids. In a 30 year-
both better timber quality and growth rate with great efficiency long study of polyploids from Pinus sylvestris L., Pinus contorta Doug-
(Niemisto ” et al., 2017). las, Picea abies (L.) H. Karst. and Larix sibirica Ledeb. in Sweden
Recent studies have sought to specifically characterize the correla- (Johnsson, 1975), the trees remained tetraploid but cone production did
tion between ploidy levels and lignin contents in trees as wood with not occur until 30 years of age. The pollen formation was abnormal,
lower lignin content is a desirable trait for certain applications. In preventing generation of triploids and seed quality was very poor in the
Arabidopsis, induced tetraploid plants producing higher biomass than C1 generation generated from backcrosses to the diploid parents. As the
the diploids also had a lower lignin content (Corneillie et al., 2019). polyploids displayed growth delays and even dwarf growth, further ef-
However in wild-type Populus tremula x P. alba, the tetraploid plants had forts towards applying polyploids to these breeding programs were
lower biomass production and a higher lignin content than the diploids discouraged (Ahuja, 2005).
(Wouters et al., 2022), whereas in a synthetic poplar allotriploid induced
by Populus pseudosimonii, P. nigra ‘Zheyin3#’ and P. beijingensis hybrid- 9. Perspectives
ization, the lignin content was reduced compared to the diploids (Xu
et al., 2022). A lower lignin content in triploids was also found in Populus Forestry will play a significant role in regaining control of the
x tomentosa Carri%ere (Yao and Pu, 1998) and Salix hybrids (Serapiglia terrestrial carbon cycle from the over-exploitation of fossil fuels, and
et al., 2015). The mechanism suggests a regulatory function for lignin polyploid trees could play an important role by accelerating biomass
biosynthesis associated with triploids. accumulation. For most forestry tree species, however, there is no data
In fruit trees, artificial polyploids are sometimes characterized by on the effects of increased ploidy on fitness. Increased knowledge of the
dwarfism (Ruiz et al., 2020). However, field trials of important forestry potential benefits of elevated ploidy levels from either allo- or auto-
species, including Acacia senegal (Diallo et al., 2016), Liriodendron x polyploidization would directly benefit forestry and guide future
sino-americanum P.C.Yieh ex C.B Shang & Z.R.Wang (Chen et al., 2021) research in this area. As we have reviewed above, increased ploidy may
and Populus tremula (P” arnik et al., 2014) show increased growth in in some cases have positive outcomes on traits like growth, biomass,
polyploids compared to diploids. In a short term experiment, triploid timber properties and resistance to climate related stressors.
hybrid Populus clones grew faster than diploids and tetraploids formed To address the knowledge gap in chromosome counts for plants,
from the same parental origins (Zhao et al., 2015; Zhang et al., 2019b). particularly trees, focused efforts in chromosome counting or flow
However, there was no unidirectional response of growth upon poly- cytometry are needed, especially in biodiversity hotspots such as Brazil,
ploidization in Salix viminalis L., where clones of autotetraploids Southeast Asia, and Madagascar. Targeting selected forest tree genera
exhibited either faster or slower growth compared to the parental for comprehensive chromosome counting and obtaining intraspecific
diploid (Dudits et al., 2016). Similar results were obtained in Populus chromosome numbers will aid in conservation efforts and help identify
hybrids where, depending on the line, polyploids generated by proto- biodiversity (Severns & Liston, 2008).
plast fusion showed both superior and inferior biomass compared to the While numerous studies indicate that polyploidy leads to larger
parental lines (Hennig et al., 2015). Triploid tree species such as Chinese cellular structures at the leaf surface, there is little knowledge on how
white poplar (Populus x tomentosa) and willow (Salix spp.) have been the structural development inside leaves, stems and roots are affected.
used as model tree species to explore growth traits. For these species, This also means that links between anatomy and physiology are poorly
triploids had increased growth and a more desirable wood composition, understood for polyploids. In particular, the ratio between photosyn-
which can be exploited in breeding programs aimed at optimizing bio- thesis and transpiration (water use efficiency) may be increased
fuel production (Zhang et al., 2012; 2013; Serapiglia et al., 2015). following polyploidization, but there is a large gap in data despite it
Other angiosperm trees with high commercial value have also been being easily measurable using for example carbon isotopic discrimina-
shown to develop better traits when growing as artificial polyploids. tion. The often hypothesized superior performance of polyploids under
Paulownia Siebold & Succ. are adaptable and fast-growing deciduous high CO2 environments has received surprisingly little attention by
trees that provide raw material for manufacturing and agroforestry, and physiologists and it should be a priority to test this, especially when
many traits related to growth and stress resistance were found to be combined with stress trials.
superior in the colchicine induced autotetraploid Paulownia australis Along with understanding the ecophysiological significance of
(PA4) compared to the parental diploid (PA2) (Xu et al., 2015). polyploidy, experiments on the polyploidization of forest trees can
Globally, Eucalyptus is the most important angiosperm tree genus for further clarify the molecular basis for the control that genome size exerts
pulp production (Myburg et al., 2014). Efforts to improve wood traits on plant form, anatomy and physiology. Experimental polyploidization
through polyploidization have shown promising results for eucalypt of key forest trees may also refine inferences on the paleo record espe-
hybrids (E. grandis x E. urophylla) where the wood from synthetic cially for stomatal density/index relationships, since induced ploidal
polyploid clones had lower basic density and longer fibers compared to series could provide contemporaneous reference collections and extend
diploids (da Silva Souza et al., 2021). The two parental genomes were data observations in the fossil record from the nearest living equivalents.
also shown to differ significantly in the repertoire of duplicated genes A close monitoring of new polyploids under field conditions is necessary,
and for structural variation (Shen et al., 2023), highlighting how allo- not only to evaluate the performance outside laboratories and green-
polyploidization can benefit forestry related trait breeding through the houses, but also to evaluate the potential invasiveness if polyploids
combination of significant genome diversity. Artificially induced poly- prove superior to conspecific diploids.
ploids have also been obtained for E. dunnii as part of the breeding Taken together, polyploidy provides substantial potential for
program to obtain new genotypes with improved adaptability and breeding of both horticultural and forestry trees for traits facilitating
possibly reduced flowering age (Castillo et al., 2020). However, in adaptation to the changing climate, yet this potential needs to be studied
another study of E. grandis x E. urophylla hybrids, polyploids exhibited on a case-by-case basis in different tree species.
decreased growth compared to their diploid counterparts (da Silva
Souza et al., 2021). Similar results were obtained in a study comparing CRediT authorship contribution statement
polyploid and diploid E. polybractea R.T.Baker grown in field and
greenhouse conditions where the polyploids were shorter and thinner Jensen Anna Monrad: Writing – review & editing, Writing –

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A. Ræbild et al. Forest Ecology and Management 560 (2024) 121767

original draft. Immanen Juha: Writing – review & editing, Writing – 2023 (Excel file).
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