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Silva Fennica 1926-1997
Acta Forestalia Fennica

Articles by Markku Nygren

Category: Article

article id 5598, category Article
Timo Kuuluvainen, Kari Leinonen, Markku Nygren, Antti Penttinen. (1996). Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests: an example from boreal spruce forest in southern Finland. Silva Fennica vol. 30 no. 2–3 article id 5598.
Keywords: boreal forests; Norway spruce; biodiversity; Picea abies; stand structure; semivariance; spatial analysis; K-function; structural variation
Abstract | View details | Full text in PDF | Author Info

The horizontal and vertical stand structure of living trees was examined in a managed and in a primeval Norway spruce-dominated forest in Southern Finland. Tree size distributions (DBHs, tree height) were compared using frequency histograms. The vertical distribution of tree heights was illustrated as tree height plots and quantified as the tree height diversity (THD) using the Shannon-Weaver formula. The horizontal spatial pattern of trees was described with stem maps and quantified with Ripley's K-function. The spatial autocorrelation of tree sizes was examined with semivariogram analysis. In the managed forest the DBH and height distributions of trees were bimodal, indicating a two-layered vertical structure with a single dominant tree layer and abundant regeneration in the understory. The primeval forest had a much higher total number of trees which were rather evenly distributed in different diameter and tree height classes. The K-function summaries for trees taller than 15 m indicated that the primeval stand was close to complete random pattern. The managed stand was regular at small distances (up to 4 m). The semivariograms of tree sizes (DBH tree height) showed that the managed forest had a clear spatial dependence in tree sizes up to inter-tree distances of about 12 meters. In contrast, the primeval spruce forest had a variance peak at very short inter-tree distances (< 1 m) and only weak spatial autocorrelation at short inter-tree distances (1–5 m). Excluding the understory trees (h < 15 m) from the analysis drastically changed the spatial structure of the forest as revealed by semivariograms. ln general, the structure of the primeval forest was both horizontally and vertically more variable and heterogeneous compared to the managed forest. The applicability of the used methods in describing fine-scale forest structure i discussed.

  • Kuuluvainen, E-mail: tk@mm.unknown (email)
  • Leinonen, E-mail: kl@mm.unknown
  • Nygren, E-mail: mn@mm.unknown
  • Penttinen, E-mail: ap@mm.unknown
article id 5410, category Article
Markku Nygren. (1990). Männyn ja kuusen siementen massan vaihtelusta. Silva Fennica vol. 24 no. 1 article id 5410.
English title: Variation in the seed mass of Scots pine and Norway spruce.
Original keywords: kuusi; mänty; lämpötila; sademäärä; siementen paino; siementen kypsyminen
English keywords: Pinus sylvestris; Picea abies; precipitation; temperature; seed mass; seed ripening
Abstract | View details | Full text in PDF | Author Info

Seed mass within any plant species is one of the least plastic components of plant structure. The aim of this study was to analyse the variation in the seed mass of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.) in relation to three environmental factors: soil fertility, mean temperature and precipitation during seed filling period. Data published earlier on seed mass of these species on different sites and different years was used in the study.

The seed mass of both species was independent of soil fertility (forest type) but did vary between different years. It is hypothesized that if the seed-ripening summer is warmer than average, Scots pine seed mass tends to be smaller. In this study, seed mass varied independently of the amount of precipitation during the ripening summer. However, generalization of the results requires further study.

The PDF includes an abstract in English.

  • Nygren, E-mail: mn@mm.unknown (email)
article id 5262, category Article
Markku Nygren. (1986). Männyn siementen syyskeräys. Silva Fennica vol. 20 no. 1 article id 5262.
English title: Autumn harvested Scots pine seeds: the effect of cone storage and germination conditions on germination capacity.
Original keywords: mänty; itävyys; dormanssi; siemenkeräys; lepotila; käpyjen varastointi
English keywords: Pinus sylvestris; Scots pine; germination capacity; seed dormancy; cone storage
Abstract | View details | Full text in PDF | Author Info

Six seed collections were made in September–December 1984 in a natural Scots pine (Pinus sylvestris L.) stand in Southern Finland. The seeds were germinated immediately after the cone collection and three photoperiods (0.8 and 24 hours) were used in germination tests.

The seeds collected in September and October possessed relative dormancy, i.e. they did not germinate in darkness and at 10°C. Later in November and December the seeds were capable to germinate in darkness and at low temperature also. The gradual change in germination capacity is attributed to chilling temperatures in natural environments or in cone storage.

The PDF includes a summary in English.

  • Nygren, E-mail: mn@mm.unknown (email)

Category: Article

article id 7648, category Article
Markku Nygren. (1987). Germination characteristics of autumn collected Pinus sylvestris seeds. Silva Fennica vol. 0 no. 201 article id 7648.
Keywords: Pinus sylvestris; Scots pine; seed maturation; seed germination; seed dormancy; chilling requirement
Abstract | View details | Full text in PDF | Author Info

Germination tests in varying photoperiod- and temperature-regimes showed that for early autumn collections, germination of Scots pine (Pinus sylvestris L.) seeds is delayed, especially at low incubation temperature (+10°C) and in darkness. The presence of light during germination (8- or 24-hour photoperiod) or high incubation temperature (+20°C) enhanced germination. As autumn proceeded, a greater proportion of seed were able of germinate in darkness and also in low temperature regime. This result was consistent in both populations studied – in seeds from natural stand (Hyytiälä, Southern Finland) and in seeds from the Hyytiälä clone archive trees, growing in the same site.

An attempt was made to relate the development of germinability during autumn to previously accumulated chilling unit (optimum temperature +3.5°C) sum. Germination percent variation in subsequent cone-collection could not, however, be explained with accumulated chilling.

The PDF includes a summary in Finnish.

  • Nygren, E-mail: mn@mm.unknown (email)
article id 7624, category Article
Pertti Hari, Seppo Kellomäki, Annikki Mäkelä, Pirkko Ilonen, Markku Kanninen, Eeva Korpilahti, Markku Nygren. (1982). Metsikön varhaiskehityksen dynamiikka. Silva Fennica vol. 0 no. 177 article id 7624.
English title: Dynamics of early development of tree stand.
Original keywords: tiheys; mänty; metsikön rakenne; metsikön kehitys; valo-olosuhteet; yhteyttäminen
English keywords: Pinus sylvestris; stand structure; photosynthesis; Scots pine; density; stand development; light conditions
Abstract | View details | Full text in PDF | Author Info

The report concludes a series of studies on the early development of young Scots pine (Pinus sylvestris L.) stands. The basis assumption made in the study series was that the within-stand light regime is the main driving force for total tree growth and its allocation of photosynthates for crown, stem and root growth. An individual tree growing in a stand under a varying light regime which is controlled by the stand structure, is the basic unit used in the study. The photosynthesis of an individual tree is determined by the light regime. The stand is formed from individual trees.

The model is applied in simulation of the growth and development of tree stands. Several computer runs representing various densities, height distributions and tree species mixtures were carried out. Potential application areas, properties of the model and future needs of investigations are discussed.

The PDF includes a summary in English.

  • Hari, E-mail: ph@mm.unknown (email)
  • Kellomäki, E-mail: sk@mm.unknown
  • Mäkelä, E-mail: am@mm.unknown
  • Ilonen, E-mail: pi@mm.unknown
  • Kanninen, E-mail: mk@mm.unknown
  • Korpilahti, E-mail: ek@mm.unknown
  • Nygren, E-mail: mn@mm.unknown

Category: Research article

article id 1056, category Research article
Katri Himanen, Markku Nygren. (2015). Seed soak-sorting prior to sowing affects the size and quality of 1.5-year-old containerized Picea abies seedlings. Silva Fennica vol. 49 no. 3 article id 1056.
Keywords: Norway spruce; seed treatment; seed radiography; seedling emergence; seedling morphology
Highlights: After soak-sorting all sunken seeds (bottom fraction) were full and viable, whereas floating seeds contained larvae-filled and immature seed; Seedlings originating from the bottom fraction were greater in height and diameter than control seedlings or those originating from the floating seeds; The proportion of saleable seedlings was four percentage points higher in the bottom fraction than in the other seedlings.
Abstract | Full text in HTML | Full text in PDF | Author Info

We studied the effect of soak-sorting Norway spruce (Picea abies (L.) H. Karst.) seeds on emergence, development and quality of container seedlings in two commercial seed lots. The seeds, separated by soaking into bottom and surface fractions, were sown in June, and the seedlings were grown during two growing seasons under typical Finnish nursery conditions. The first summer seedlings were grown in a greenhouse and outdoors for the second, full growing season. All sunken seeds were full and viable according to radiography, whereas the floating seeds contained 2% and 13% larvae-filled and 8% and 11% anatomically immature seeds, depending on the seed lot. Seedlings grown from the bottom fraction seed emerged 2.5–3.5 days earlier than seedlings of storage dry (i.e. control) seed. Height, diameter, and shoot and root dry mass of the seedlings were affected by soaking after both the first and second growing seasons. The largest seedlings originated from the bottom fraction. The proportion of saleable seedlings was four percentage points higher in the bottom fraction than in the other seedlings. The effects of soaking found in this study are more notable than as previously reported for Norway spruce seedlings. This suggests that soaking and soak-sorting may be most useful when the growing conditions are stressful, i.e. when seeds are sown in summer rather than 1-year-old seedling crops sown in spring under the climate conditions typical of Finland.

  • Himanen, Natural Resources Institute Finland (Luke), Green technology, Juntintie 154, FI-77600 Suonenjoki, Finland E-mail: (email)
  • Nygren, Natural Resources Institute Finland (Luke), Management and Production of Renewable Resources, Juntintie 154, FI-77600 Suonenjoki, Finland E-mail:

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