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Forest Ecology and Management 255 (2008) 2870–2875

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Ash-fertilization improves germination and early establishment of birch

(Betula pubescens Ehrh.) seedlings on a cut-away peatland
Noora Huotari, Eila Tillman-Sutela *, Jorma Pasanen, Eero Kubin
The Finnish Forest Research Institute, Muhos Research Unit, Kirkkosaarentie 7, FIN-91500 Muhos, Finland
Received 5 October 2007; received in revised form 18 January 2008; accepted 22 January 2008

Abstract
Common objectives of afforestation of cut-away peatlands are the diminution of nutrition loads and the production of energy wood in an
efficient and economic way. We studied the effect of fertilization and afforestation methods on the early establishment of downy birch (Betula
pubescens Ehrh.) seedlings on a cut-away peatland. Six treatments of peat-ash, wood-ash, biotite, or PK-fertilizer were replicated in three blocks in
different mixtures corresponding to 50 kg/ha of phosphorous. Birch seeds were sown in randomized halves of split plots, while other halves were
naturally afforested. Natural afforestation was equal to sowing in terms of seedling amounts due to closeness of surrounding birch stands with
abundant seed crops. After the first growing season the greatest quantities and dominant heights of birch seedlings appeared on the ash-fertilized
plots, whereas on the unfertilized areas both the establishment and growth of seedlings was very poor. On the other hand, the quantity of Scots pine
(Pinus sylvestris L.) seedlings was more than three times higher on the unfertilized areas compared to the ash-fertilized areas. The differences
between the fertilized and the unfertilized areas accentuated during the four observation years. Our results support regular use of ash-fertilizers for
the afforestation of cut-away peatlands instead of the ash being deposited as a waste in landfills.
# 2008 Elsevier B.V. All rights reserved.

Keywords: Afforestation; Peat-ash; Wood-ash; PK-fertilizer; Pinus sylvestris

1. Introduction reed canary grass (Phalaris arundinacea L.) cultivation, re-

paludification and afforestation (Selin, 1996). Afforestation is
Harvesting of peatlands for energy production changes the an advantageous and popular after-use option since it has both
original mire ecosystem completely and practical methods for economical and aesthetic values. The efficient ways to increase
the after-use of these disturbed ecosystems are of major the post-harvesting growth of tree stands on cut-away areas still
importance for both landscape and environment. Post-harvest- considering environmental effects are currently studied in
ing area is a harsh environment for plants due to the unstable Finland, Ireland and Sweden.
water conditions, wind erosion and frost heaving, which Downy birch (Betula pubescens Ehrh.) is a suitable endemic
hamper the establishment of plants during several decades species for afforestation of cut-away peatlands due to its ability
(Price et al., 2003; Petrone et al., 2004; Campbell et al., 2002; to grow on moist soils and the relatively short rotation time to
Groeneveld and Rochefort, 2002; Lavoie et al., 2005). In produce energy wood on fertile sites (Ferm, 1993). Propagation
addition, decreased microbial populations (Croft et al., 2001; of birch by seeds is, however, often problematic on peatlands
Andersen et al., 2006) and especially low mineral nutrient even if seeds germinate on cushions of Sphagnum, because the
contents (Salonen and Laaksonen, 1994; Huotari et al., 2007) young seedlings rarely establish due to very high mortalities
can restrict the formation of vegetation. Post-harvesting cut- during their first year (Kinnaird, 1974; Cabiaux and Devillez,
away sites have been adapted for agriculture, bird sanctuaries, 1977). Bare ground is most suitable for the establishment of
birch seedlings, particularly on places where vegetation has
been removed by burning (Cabiaux and Devillez, 1977) and
consequently competition for space has decreased. However,
* Corresponding author. Tel.: +358 102113719; fax: +358 102113701.
E-mail addresses: [email protected] (N. Huotari),
invasion by birch following burning (Atkinson, 1992) may also
[email protected] (E. Tillman-Sutela), [email protected] be due to changed nutritional conditions of the soil. On cut-
(J. Pasanen), [email protected] (E. Kubin). away peatlands the remaining peat layer is typically rich in
0378-1127/$ – see front matter # 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2008.01.062
 N. Huotari et al. / Forest Ecology and Management 255 (2008) 2870–2875 2871

organic nitrogen, but poor in mineral nutrients, especially were established on the experimental field of 3 ha and each of
phosphorus and potassium (Paavilainen and Päivänen, 1995; them was divided into six 0.16 ha trial plots. The plots were
Wind-Mulder et al., 1996). On the other hand mineral sub-soil split and 1 kg ha1 of downy birch seeds were hand-sown in
of cut-away peatland can also be poor in nitrogen, and seedlings early September 2000 on randomized halves, while the other
may consequently be stunted. For these reasons the natural halves were afforested naturally. In May 2001 the following
establishment of plants on these areas is in most cases slow and fertilization treatments were randomized between the plots
patchy. Due to its high quantities of mineral nutrients and and replicated in the three blocks: (1) unfertilized; (2) wood-
positive effect on the mineralization of organic nitrogen (Park ash 7874 kg ha1; (3) wood-ash 3937 kg ha1 + peat-ash
et al., 2005), ash may be advantageous to establish trees on cut- 2397 kg ha1; (4) peat-ash 4794 kg ha1; (5) peat-ash
away peatlands. 4794 kg ha1 + biotite 1500 kg ha1; and (6) Forest PK-
Most studies on the effects of ash-fertilization on afforesta- fertilizer 556 kg ha1. The nutrient contents of the ashes were
tion results on cut-away peatlands or peatlands drained for analyzed in the laboratory of Muhos Research Station of the
forestry have had a goal of growing conifers for sawmill Metla (see Table 1 in Huotari et al., 2007) and the doses of
industry or to produce biomass in short-rotation monocultures fertilizers per plot were adjusted corresponding to 50 kg ha1
of exotic willow (Salix spp.) species. On cut-away peatlands of phosphorus, the quantity recommended in peatlands drained
only very high application rates of ash have increased the for forestry (Paavilainen and Päivänen, 1995).
biomass production on willow plantations (Hytönen, 1998; Nine circular sample plots of 0.5 m2 (d = 0.78 m) were
Park et al., 2005). Wood-ash is known to promote tree growth systematically placed on each trial plot and the amounts and
on peatlands drained for forestry and the influence is reported to dominant heights of established seedlings were measured by
be long lasting (Silfverberg, 1996; Moilanen et al., 2002). The species yearly in 2001–2004. In this context the dominant
effect and duration of ash-fertilization may be, however, height refers to the tallest seedling of each species and age
different on cut-away peatlands where the original vegetation group in the circular sample plot.
has been completely removed and the depth of the remaining Normality of the data was tested using Shapiro–Wilkin test
peat layer varies considerably. In addition, the results from the and homogeneity of variances using Levene’s test. The effect of
effect of wood-ash on the germination of pine or birch seeds fertilization and afforestation methods on the quantity of birch
have been contradictory (Rikala and Jozefek, 1990; Reyes seedlings, which germinated in 2001, was tested using the two-
et al., 1997; Reyes and Casal, 2004), and the research on the way analysis of variance. Since there were no statistically
early development of seedlings or on the afforestation methods significant differences in seedling quantities between the two
on the ash-fertilized cut-away peatlands is very scanty (Huotari afforestation methods and no interaction between the affor-
et al., 2007). For these reasons we studied the effect of ash- estation and fertilization treatments in 2001, the data was
fertilization and afforestation methods on the germination and combined, and one-way ANOVA was then used to detect the
early establishment of downy birch seedlings on a cut-away differences in total amounts of birch seedlings among the
peatland. fertilization treatments in 2004. Tukey’s multiple comparisons
test was used to determine the differences among the
2. Materials and methods fertilization treatments. Both the data concerning the year
2001 and 2004 were square-root transformed prior to the tests.
The experimental area is located on a cut-away peatland in The effect of fertilization treatment on the dominant height
Northern Ostrobothnia (648440 N, 258160 E, 45 m a.s.l.), in of birch seedlings during 2001–2004 was analyzed using
Finland. The total area of 385 ha was vacuum-mined for Repeated Measures ANOVA. The between-subjects factor used
horticultural peat and energy generation by 1996 and the depth was fertilization treatment and the within-subjects factor time.
of the remaining peat layer on the area varied from 20 to 53 cm. Tukey’s multiple comparisons test was used to determine the
The soil below the peat was sandy till and ground water level differences among the fertilization treatments. The 5%
was about 50 cm. The residual peat was medium humified significance level was applied in all the statistical analysis.
moss-sedge peat (SC-p, H5) with some wood residues, and the
pH-value varied from 4.6 to 5.1. The nitrogen content in the 3. Results
residual peat layer was on average 2.3%, which is higher than
the N concentrations evaluated to be adequate (1.5–2.0%) for 3.1. Quantity of birch seedlings
tree growth on drained peatlands (Moilanen, 1993; Pietiläinen
and Kaunisto, 2003). The average yearly temperature sum in Fertilization had a highly significant effect on the quantity of
the area is 1040 d.d. The amount of precipitation from 1st April birch seedlings after one growing season (F = 47.524, d.f. = 5,
to 15th July in 2001 was 236 mm, in 2002 76 mm, in 2003 p = 0.000), whereas the differences in the seedling quantities
116 mm and in 2004 194 mm. Birch stands of 8–12 m between the two afforestation methods (F = 3.565, d.f. = 1,
(dominant height) surrounded two sides of the area at a p = 0.060) or interaction between the afforestation and
distance of 100–150 m. fertilization methods in 2001 (F = 1.046, d.f. = 5, p = 0.391)
The cut-away area was drained in August 2000 by cleaning were confirmed as statistically non-significant. Due to these
the old ditches at 40 m distance but no other soil preparation reasons and to the fact that sowing was conducted only in 2000,
was done on the non-vegetated cut-away area. Three blocks whereas there was some natural dissemination also in ensuing
2872 N. Huotari et al. / Forest Ecology and Management 255 (2008) 2870–2875

Fig. 1. The quantities of birch seedlings in 2001 on the treatments: No = un- Fig. 2. Total quantities of birch seedlings of varying ages observed in 2004 on
fertilized, Wa = wood-ash, Ma = mixed-ash, Pa = peat-ash, Pa + B = peat- the six treatments: No = unfertilized, Wa = wood-ash, Ma = mixed-ash,
ash + biotite, PK = forest PK-fertilizer. The data of the two afforestation Pa = peat-ash, Pa + B = peat-ash + biotite, PK = forest PK-fertilizer. The data
methods is combined. Bars represent means and vertical lines standard errors. of the two afforestation methods is combined. The total seedling quantities
The seedling quantities indicated with different letters differ from each other at indicated with different letters differ from each other at a 5% level of
a 5% level of significance. significance.

years, the data of the two afforestation treatments was L.) seedlings was almost 8000 plants per hectare on the
combined and the results presented hereafter concern only unfertilized plots, whereas on ash-fertilized areas their amounts
the effects of fertilization. were notably less, from 0 to 1500 plants per hectare. On the
Seedling establishment after one growing season was on all Forest PK-fertilized plots there were almost 3000 pine
fertilized plots superior to that on the unfertilized plots, where seedlings per hectare, germinated in 2001, and the amount
the number of birch seedlings was less than 16,000 plants per of seedlings stayed rather stable during the observation period.
hectare (Fig. 1). The quantity of birch seedlings in the ash-
fertilized areas varied from 236,000 to 267,000 plants per 3.2. The dominant height of seedlings
hectare, whereas 108,000 plants per hectare appeared on the
plots where Forest PK-fertilizer had been used. The difference Fertilization had a significant effect on the dominant heights
between the Forest PK-fertilized area and all the ash-fertilized of birch seedlings across the 4 years observation period
areas were also statistically significant. (time  fertilization effect F = 8.364, d.f. = 7.898, p = 0.000).
Three years later more than 80% of the birch seedlings, In 2001 the greatest dominant heights, 9 cm, were measured on
which had germinated in 2001 were still alive on the fertilized the plots that were fertilized with the mixture of peat- and
plots and the total amount of seedlings had even increased due
to natural dissemination (Fig. 2). Meanwhile, the number of
original birch seedlings on the unfertilized plots had decreased
drastically and only 14% of the seedlings germinated in 2001
were alive. Even if new seedlings started due to natural
dissemination during the four observation years, there were
only a few plants that survived on the unfertilized plots after
their germination year.
Seedlings of other tree species also appeared on the trial
plots due to natural dissemination from the surrounding area.
The highest amount of willow (Salix spp.) seedlings, about
80,000 plants per hectare, was established during the first
growing season on the plots, where peat-ash or the mixture of
wood- and peat-ash was used (not shown here). A somewhat
smaller number, 50,000–60,000 seedlings per hectare grew on
the plots where wood-ash, Forest PK-fertilizer or peat-ash
mixed with biotite was used, whereas only 4000 seedlings per
Fig. 3. Annual growth in dominant heights of birch seedlings germinated in
hectare germinated on the unfertilized plots. The number of
2001 on the fertilization treatments. The data of the two afforestation methods
willow seedlings had remained rather stable on the fertilized are combined. Fertilization treatments: No = unfertilized, Wa = wood-ash,
plots after four growing seasons. Distinguishing from birch and Ma = mixed-ash, Pa = peat-ash, Pa + B = peat-ash + biotite, PK = forest PK-
willow seedlings, the quantity of Scots pine (Pinus sylvestris fertilizer.
 N. Huotari et al. / Forest Ecology and Management 255 (2008) 2870–2875 2873

wood-ash (Fig. 3). The use of pure peat-ash also resulted in the mineral soil have been mixed together (Lumme, 1988). In
good growth of birch seedlings, the dominant height being 7 cm these cases the main reason for no marked effect of the ash
after the first growing season. In the plots where other fertilizers application on the survival of birch seedlings, may be that the
were used the dominant heights were slightly smaller, 4–5 cm, mineral nutrient condition of the soil was already adequate for
whereas on the unfertilized plots the dominant heights were no the establishment of tree seedlings. The effect of ash on plant
more than 2 cm. germination is found to vary in magnitude depending on the
In 2004 the greatest dominant heights, 67 cm, were on the family and species (Gonzáles-Rabanal and Casal, 1995). The
plots where the mixture of peat- and wood-ash had been used, absence of pine seedlings on the ash-fertilized plots but
and the superiority of this mixture to other fertilizers except to occurrence on the unfertilized plots confirmed the reported
pure peat-ash was also confirmed as statistically significant retarding effect of ash on germination of pine seeds (Reyes and
(Fig. 3). The seedlings growing on peat-ash fertilized areas had Casal, 2004).
dominant heights of 58 cm, and the dominant heights of birch The difference in the amounts of live birch seedlings,
seedlings on other fertilized plots varied from 46 to 53 cm. The germinated in 2001, among the fertilized and unfertilized areas
birch seedlings on the unfertilized areas were clearly smaller, was emphasized during the four observation years: in 2004 the
with dominant heights of about 11 cm after four growing quantity of original birch seedlings on the unfertilized plots was
seasons and the difference compared to the fertilized areas was only 14% of the first year’s amount. Our results thus confirmed
also statistically significant. Moreover, the seedlings on the the earlier observations on the high mortality of young birch
unfertilized areas had an anomalous curly appearance. seedlings on harvested peat surfaces (Salonen et al., 1992).
The dominant heights of naturally established willow (Salix High post-germination mortality typical for birch seedlings has
spp.) seedlings after the first growing season were largest, from frequently been interpreted as a result of drought (Kinnaird,
8 to 10 cm on the plots where ash-fertilizers had been used. 1974; Cabiaux and Devillez, 1977; Bérubé and Lavoie, 2000).
Forest PK-fertilizer resulted in slightly smaller heights, 6 cm, The high and almost unchanged quantities of birch seedlings on
whereas on the unfertilized areas the dominant heights of all fertilized plots proved that the post-germination survival was
willow seedlings were clearly smaller, 2 cm. The later height also highly dependent on the nutrition level of the soil. This
development of the willow seedlings continued as during the conclusion is also in accordance with the morphology of birch
first year without notable differences between the fertilization seeds, which have very small endosperm as a nutrient reserve
treatments. After four growing seasons the dominant heights of and therefore cannot survive a long time after germination on
the willow seedlings germinated in 2001 varied on the ash- cushions of Sphagnum or on unfertile bare ground.
fertilized plots from 38 to 47 cm. On the PK fertilized areas the In 2004, the total amounts of birch seedlings of varying ages
dominant height was 31 cm and on the unfertilized areas 11 cm. were still almost two times higher on the ash-fertilized plots
On the other hand, the dominant heights of Scots pine compared to the Forest PK-fertilized plots, in spite of the
seedlings germinated in 2001 were 3–4 cm after the first natural birch dissemination during the four observation years.
growing season and there were no major differences between This result indicates that ash-fertilizers were more appropriate
the fertilization treatments or between the fertilized and to the establishment and survival of birch seedlings than the
unfertilized plots. After four growing seasons pine seedlings on commercial Forest PK-fertilizer on a cut-away peatland. The
the areas fertilized using peat-ash reached dominant heights of nitrogen in cut-away peatlands is mainly organically bound in
72 cm, whereas the dominant heights on the other ash-fertilized the peat and not usable for plants as such, whereas ash-
areas varied from 41 to 55 cm. Thus, they had about the same fertilization is reported to accelerate the conversion of
dominant heights as birch seedlings on ash-fertilized plots. nitrogenous compounds in peat into a form that can be utilized
Dominant heights of pine seedlings on the unfertilized or PK- by plants and bacteria (Karsisto, 1979; Lehtonen and Tikkanen,
fertilized plots were only slightly smaller, 38 cm, after four 1986; Hytönen, 1998). In addition, next to the enhanced
growing periods. establishment of birch seedlings, ash-fertilization is also
reported to accelerate the formation of pioneering moss cover
4. Discussion (Huotari et al., 2007). Moss cover stabilizes loose peat surface
and prevents the frost-heaving damages of seedlings, which is
Our results proved that ash-fertilization in particular reported to complicate the rooting and survival of young
improved germination and early establishment of downy birch seedlings on cut-away peatlands (Campbell et al., 2002;
seedlings on a cut-away peatland. The quantity of birch Groeneveld and Rochefort, 2002; Lavoie et al., 2005). The
seedlings in the plots where ash-based fertilizers were applied, positive impacts of ash-fertilization on tree growth are reported
was at its best 16 times greater than on the unfertilized areas and to last up to 50 years or more (Mikola, 1975; Moilanen et al.,
two times greater than on the Forest PK-fertilized areas already 2002, 2005), which is an adequately long period when
after the first growing season. This result was even more considering the energy-wood cultivation of birch with 20–30
prominent than those obtained using wood-ash on nutrient-poor year’s rotation time.
peatlands (Silfverberg, 1995). Our result, however, contra- Our results showed no major differences in the amounts of
dicted the reported effect of wood-ash on the birch establish- birch seedlings between the sown and the naturally regenerated
ment on mineral soils (Reyes et al., 1997), on abandoned fields areas. This is partly due to the closeness of the birch stands and
(Karlsson, 1996) or on a cut-away peatland, where the peat and their good seed crops, which assured a maximum density of
2874 N. Huotari et al. / Forest Ecology and Management 255 (2008) 2870–2875

young seedlings on the area observed also in earlier studies on 2006. Vapo Co and Tekes (National Technology Agency of
afforestation of low-productive peatlands (Sundström and Finland) supported this study financially. Mr. J. Issakainen
Hånell, 1999). Sufficient precipitation by mid-July in 2001 also organized the field trials, Ms. I Murtovaara finished the figures,
favored seed germination, since moist environmental condi- Dr. A. Kauppi gave valuable comments on the manuscript and
tions in early summer are known to have a major impact on the Mr. R. Gear revised the English text. We thank sincerely all who
germination and successful establishment of young seedlings have contributed to this study.
on mineral soils (de Chantal, 2003). In addition, the pH-value of
the peat was within the reported optimum for both birch and
pine seed germination (Rikala and Jozefek, 1990). References
Fertilization had a major impact on the dominant heights of
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