effects of environmental factors on grain yield and quality of oats (avena sativa ...

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This article was downloaded by: [The University of Manchester Library] On: 21 November 2014, At: 06:27 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Acta Agriculturae Scandinavica, Section B — Soil & Plant Science Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/sagb20 Effects of environmental factors on grain yield and quality of oats (Avena sativa L.) cultivated in Finland Marketta Saastamoinen a a Boreal Plant Breeding , Myllytie 10, Jokioinen, FI31600, Finland Published online: 01 Oct 2008. To cite this article: Marketta Saastamoinen (1998) Effects of environmental factors on grain yield and quality of oats (Avena sativa L.) cultivated in Finland, Acta Agriculturae Scandinavica, Section B — Soil & Plant Science, 48:3, 129-137, DOI: 10.1080/09064719809362490 To link to this article: http://dx.doi.org/10.1080/09064719809362490 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions

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This article was downloaded by: [The University of Manchester Library]On: 21 November 2014, At: 06:27Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: MortimerHouse, 37-41 Mortimer Street, London W1T 3JH, UK

Acta Agriculturae Scandinavica, Section B — Soil& Plant SciencePublication details, including instructions for authors and subscriptioninformation:http://www.tandfonline.com/loi/sagb20

Effects of environmental factors on grain yieldand quality of oats (Avena sativa L.) cultivated inFinlandMarketta Saastamoinen aa Boreal Plant Breeding , Myllytie 10, Jokioinen, FI‐31600, FinlandPublished online: 01 Oct 2008.

To cite this article: Marketta Saastamoinen (1998) Effects of environmental factors on grain yield and quality ofoats (Avena sativa L.) cultivated in Finland, Acta Agriculturae Scandinavica, Section B — Soil & Plant Science, 48:3,129-137, DOI: 10.1080/09064719809362490

To link to this article: http://dx.doi.org/10.1080/09064719809362490

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”)contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensorsmake no representations or warranties whatsoever as to the accuracy, completeness, or suitabilityfor any purpose of the Content. Any opinions and views expressed in this publication are the opinionsand views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy ofthe Content should not be relied upon and should be independently verified with primary sourcesof information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings,demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arisingdirectly or indirectly in connection with, in relation to or arising out of the use of the Content.

This article may be used for research, teaching, and private study purposes. Any substantial orsystematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distributionin any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found athttp://www.tandfonline.com/page/terms-and-conditions

Acta Agric. Scand., Sect. B, Soil and Plant Sei. 1998: 48, 129-137

Printed in Ireland. All rights reservedACTA AGRICULTURE

SCANDINAVICA

ISSN 0906-4710

Effects of Environmental Factors onGrain Yield and Quality of Oats (Avenasativa L.) Cultivated in Finland

Saastamoinen, M. (Boreal Plant Breeding, Myllytie 10, FI-31600Jokioinen, Finland). Effects of environmental factors on grain yield andquality of oats (Avena sativa L.) cultivated in Finland. Accepted Octo-ber 7, 1998. Acta Agric. Scand., Sect. B, Soil and Plant Sci. 48:129-137, 1998. © 1998 Scandinavian University Press.

Grain yield and yield quality characteristics were studied in the twooat varieties, Puhti and Veli, during 4 different years, 1987-90. Thegrowing period of 1987 was extremely cold, while 1988 was warm anddry. Oat trials were established at eight locations throughout the oatcultivation area of Finland. Information on climatic and edaphic fac-tors, as well as fertilization levels, was used to explain the variation ingrain yield and quality characteristics by multiple step-wise regressionanalysis. High temperatures during the growing period (x) signifi-cantly decreased grain yields (y) in the early variety Veli (y = 9241 —335x). Ρ fertilization was the main yield increasing factor in the laterPuhti variety. The early variety, Veli, is better adapted to cultivationin a cool climate than the later variety, Puhti. Protein content wassignificantly decreased by heavy precipitation and oil content signifi-cantly increased by a low temperature during the growing period inboth test varieties. High precipitation significantly decreased the hec-tolitre weight of both varieties. High Ρ fertilization was the mainsignificant factor in decreasing the 1000 grain weight.

Marketta SaastamoinenBoreal Plant Breeding, Myllytie 10,FI-31600 Jokioinen, Finland

Key words: climate, oil content,precipitation, protein content,temperature.

Introduction

It has been postulated that elevated global CO2 con-centration eventually increase the average global tem-perature. The impact of climatic change alsoinfluences vegetation. In the northern climate lowtemperatures and a short growing period are growth-limiting factors. According to various studies higherCO 2 concentration affects the growth of differentplant species. In the review article by Poorter (1993),concerning 156 plant species, the average growthstimulation is estimated as 37% when the atmosphericCO2 concentration is doubled.

A higher yearly mean temperature is commonlybelieved to increase the grain yield in cool northernregions. However, the optimum temperature duringthe growing period differs for various plant species.Sœbo & Mortensen (1996) found increased dry mat-ter accumulation only in Festuca rubra out of sevengrass species cultivated in a higher CO2 concentration

in a cool oceanic maritime climate.Oats have adapted to cultivation in cool northern

climates and are generally better adapted to growthin the cooler microclimate of acid peat soils than iswheat, rye or barley. Oats are chiefly cultivated innorthern regions and are not such an importantspecies further south.

Grain yield and yield quality characteristics wereexamined in two oat varieties used as standard vari-eties in oat plant breeding trials during 4 differentyears in Finland. The effects of climatic factors,average temperature of the growing period and pre-cipitation, as well as other environmental factors,were studied in the grain yields and yield quality ofthe test varieties.

Materials and methods

Grain yields, other agronomic traits, chemical qual-

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Table 1. Soil types and means for nitrogen (N), phosphorus (P) and potassium (K) fertilization levels, mean temperature of growing period and precipitation »from sowing to yellow ripening and pH, Ca, Κ, Ρ and Mg levels of the soil for different trial locations £•

οFertilization Nutrient levels of soil 5"(kg ha"1) (mg I"1) s

Temperature PrecipitationNumber of of growing of growing

Location/situation Trial trials, η Years Soil types Ν Ρ Κ Soil pH Ca Κ Ρ Mg period (CC) period (mm)

Institute of Plant Breeding, I 4 1987-90 Mould 78 31 60 5.5 3425 171 5.8 429 14.4 225JokioinenLong. 23°29' E II 4 1987-90 Silty clay, sandy clay 105 42 80 6.0 1969 278 14.4 613 14.3 180Lat. 60°49' N and fine sand

III 4 1987-90 Silty clay, sandy clay 105 42 80 6.0 1969 278 14.4 613 14.2 184and fine sand

IV 4 1987-90 Fine sand and sandy 101 40 77 6.1 1763 209 14.6 400 14.1 184clay

V 4 1987-90 Fine sand, sandy clay 101 40 77 6.1 1863 183 9.3 445 14.1 184and coarse sand

South-West Finland Res. Sta. 3 1988-90 Gyttja clay and gyttja 100 30 30 5.2 964 220 9.9 151 15.4 181Long. 21°5V E fine sandLat. 60°38' NSatakunta Res. Sta. 4 1987-90 Silty fine sand, silty 94 34 65 6.1 1527 148 13.1 222 14.1 233Long. 22°14' E clay and clayey fineLat. 61°17' N sandAnjalankoski Res. Sta. 4 1987-90 Silty clay 98 25 48 6.5 2531 245 9.4 334 15.6 173Long. 26°48' ELat. 60°43' NCentral Finland Res. Sta. 4 1987-90 Fine sand, gyttja and 70 28 54 5.1 751 80 7.0 55 13.6 298Long. 26°00' E siltLat. 62°00' NKarelia Res. Sta. 4 1987-90 Ligno Carex peat 61 24 46 4.8 1152 77 5.9 81 14.3 231Long. 30°19' ELat. 62°14' NSouth Ostrobothnia Res. Sta. 4 1987-90 Silt, clayey silt, gyttja 83 34 63 4.6 349 192 18.8 23 13.6 226Long. 22°30' E clay and fine sandLat. 62°56' N

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Grain yield and quality of oats

ity, protein and oil contents of the yields of twoFinnish oat varieties, Puhti and Veli, were deter-mined at eight different locations in Finland during4 consecutive years (1987-90). Puhti and Veli werethe standard varieties in the variety trials, compris-ing 12-42 oat varieties or lines. The experimentaloat varieties were tested in yearly trials (Table 1).Trial locations comprised the total oat cultivationarea of Finland. The trials were established accord-ing to the experimental designs of Cochran & Cox(1960). There were three or four replications in thetrials and the plot size varied from 10.00 to 13.75m2 at different locations. Fertilization was deter-mined yearly according to the soil type and nutrientlevels of the soil. Fertilization was carried out inthe spring before sowing. Soil calcium, potassium,phosphorus and magnesium contents were deter-mined according to Vuorinen & Mäkinen (1955)and expressed as mg l" 1 (Kurki et al., 1965). Thesoil types were representative of all typical soilsfound in Finland: silt, clay, gyttja, sand and or-ganic soils, mould and peat. Soil pH ranged fromvery acid (pH 4.6) to good soil pH of 6.8. Averagetemperatures and precipitation levels of the growingperiods were calculated for the test varieties fromsowing to yellow ripening. Average temperaturesand precipitation levels of the growing periods dif-fered in each experimental year. The difference be-tween the average temperature of the growingperiods for all years was 5.0°C and for precipitationwas 166 mm (Table 1).

Protein content was determined in whole milledgrains by the NIR method, using a Technicon In-fralyzer 400. The NIR instrument was calibrated bythe Kjeldahl method with Tecator Kjeltec System IIequipment. Oil content was analysed by the nuclearmagnetic resonance (NMR) method using a New-port Analyser Magnet Type 10 instrument as de-scribed previously by Saastamoinen et al. (1989).

Grain yields of the varieties were calculated ac-cording to Cochran & Cox (1960) using the SPSSxstatistical program (SPSS, 1986). The effects of en-vironmental factors on the grain yield synthesis andchemical compositions of the experimental oat vari-eties were studied by step-wise regression analysis(SPSS, 1986).

Results and discussion

Grain yield

Grain yield varied considerably during the experi-mental period. The highest yields were obtained in1990 and the lowest in 1988 for both experimentalcultivars (Table 2). The average temperature of the

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M. Saastamoinen

growing period in 1990 was 13.4°C, and in 1988 itwas 16.6°C. 1988 was the driest year.

The correlation coefficient between the grain yieldand mean temperature of the growing period wassignificant for the Veli variety (Table 3). The grainyield of the Veli variety was found to be dependenton the average temperature of the growing period(Table 4). A lower temperature during the growingperiod increased the grain yield, despite one very coldsummer, in 1987. The average temperature of thegrowing period was the only significant factor ex-plaining the variation in the grain yield of the Velivariety. Beringer (1967) reported a 30-40% increasein the grain yield of oats when the temperature of thegrowing period was low (12°C) compared with a hightemperature (30°C). Hellewell et al. (1996) found 87%higher yields in oats with 15°C day/15°C night condi-tions compared with 31°C day/15°C night conditionsin a growth-chamber study, and 12% higher yieldswith 15°C day/15°C night conditions compared with23°C day/15°C night conditions.

Environmental factors had different effects on thePuhti cultivar (Tables 3 and 4). Grain yield wasfound to be dependent on soil phosphorus and potas-sium levels. Phosphorus increased the grain yield andpotassium decreased it. The correlation coefficientbetween grain yield and mean temperature of thegrowing period was negative in the case of the Puhtivariety, but temperature was not a significant factorexplaining the yield level. The above-mentioned envi-ronmental responses can perhaps be explained by thedifferent growing times and general adaptation of thetest varieties. Veli is 5 days earlier than Puhti andproduces higher yields in cold years in comparisonwith late varieties. Of the two test cultivars, Veli isbetter adapted to a cool climate. Hellewell et al.(1996) found higher yields in early, mid-early andlate oat varieties with 15°C day/15°C night than with31°C day/15°C night temperature. In this study theaverage growing temperature was much lower in thecold year of 1987.

The results suggest that the optimum temperaturefor the cultivation of oats is not very high. Consider-able research has been carried out on the differentoptimum growth temperatures for cool-season (C3)and warm-season (C4) plant species. There is a greatdifference between the C 3 and C 4 plant species interms of the optimum temperature for photosynthe-sis. C 4 species can maintain photosynthesis at highertemperatures than C 3 species (Pearcy & Ehleringer,1984). However, C4 species have a higher capacity toutilize water (Stout, 1992) and soil N (Brown, 1985).There are few studies on the different optimum tem-peratures for growth in different C 3 species and geno-types. The present results indicate that the optimum

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Table 3. Correlation coefficients (r) between different agronomie and quality characteristics and the environmental factors in Veli and Puhti varieties

CO

ω

Variety/characteristics

Veli (n = 47)Grain yieldGrowing timePlant heightLodging1000 grain sizeHl-weightProtein contentHull contentOil content

Puhti {n =47)Grain yieldGrowing timePlant heightLodging1000 grain sizeHl-weightProtein contentHull contentOil content

*P<0.05; * * P < 0

Mean temperature

-0.45**-0.96***-0.66***-0.42**-0.17

0.48***0.45**0.31*

-0.69***

-0.20-0.96***-0.67***-0.38**

0.250.58***0.51***

-0.03-0.78***

.01; ***P<0.001.

Precipitation

0.170.80***0.54***0.53***

-0.25-0.57***-0.68***-0.23

0.57***

-0.120.82***0.53***0.41**

-0.41**- 0 . 6 1 * * *-0.70***-0.01

0.67***

Fertilization

N

-0.24-0.20-0.44**-0.04-0.05-0.05

0.260.38**

-0.09

-0.20-0.21-0.43***-0.07-0.13

0.010.200.32*

-0.05

Ρ

-0.170.09

-0.080.19

-0.35*-0.32*-0.05

0.39**0.09

-0.34*0.13

-0.060.09

-0.42**-0.27-0.10

0.32*0.20

Κ

-0.120.11

-0.060.11

-0.24-0.25-0.04

0.38**0.06

-0.230.11

-0.040.03

-0.28-0.20-0.03

0.33*0.13

Soil mineral

Κ

-0.28-0.23-0.41**-0.01-0.06-0.05

0.33*0.27

-0.06

-0.23-0.22-0.37**-0.11-0.09

0.020.29*0.05

-0.03

content

Ρ

0.160.070.05

-0.120.280.290.15

-0.31*-0.18

0.34*0.030.10

-0.020.230.240.10

-0.35*-0.15

Mg

-0.09-0.08-0.24

0.18-0.05-0.31*

0.050.35*0.12

-0.25-0.04-0.24-0.04-0.21-0.21

0.050.41**0.16

Ca

0.040.00

-0.020.250.15

-0.250.140.110.08

0.000.01

-0.010.100.03

-0.200.090.250.06

- pH ofsoil

-0.030.01

-0.170.100.19

-0.110.020.04

-0.02

0.01-0.01-0.15-0.00

0.10-0.07-0.05

0.150.01

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Table 4. Dependence of grain yield (y) on mean temperature of growing period (χΊ), phosphorus (x2) andpotassium (x3) level of the soil in Veli and Puhti varieties in the years 1987-90 analysed by multiple step-wiseregression analysis

Variety

VeliPuhti

Numberof trials

4747

Equation

y= 9240.95-335y = 4996.87 + 51

.17χη

.59x2-6.05x3

r2 100

44.6649.84

(%)F-Value forequation

11.21**7.27**

r-Value

-3.35**

for:

x2

3.39** -2.77**

Constant

6.45***12.18***

** P<0.01; *** P<0.001.

temperature may differ for different oat genotypes,because the average temperature of the growing pe-riod was significant in explaining the grain yields ofthe Veli variety but not those of Puhti. Yields of Veliwere significantly higher at lower growth tempera-tures. The results are consistent with those of Mela etal. (1996), in that higher yields can be obtained in awarmer climate only by means of better adaptedgenotypes.

Protein content

Protein synthesis was likewise shown to be dependenton environmental variation. Significant positive cor-relation coefficients were found between protein con-tent and the average temperature of the growingperiod, and significant negative correlations betweenprecipitation and the protein content of both testvarieties (Table 3). The protein content of the twovarieties, Puhti and Veli, were dependent on precipi-tation levels calculated by multiple step-wise regres-sion analysis (Table 5). A high precipitation levelduring the growing period caused a decrease inprotein content. Average temperature was not, how-ever, a significant factor explaining the variation inprotein content. High precipitation increases leach-ing, leaving plants with less available nitrogen in wetconditions.

The effect of drought in increasing protein contentis very well known, especially in barley and wheat(Schwanitz & Schwarze, 1937a,b; Postel, 1956; Sow-ers et al., 1994). Nitrogen fertilization did not in-crease protein content because the fertilization level

was adjusted according to the level of nutrients in thesoils. In previous studies it has been found thatnitrogen fertilization increases the protein content ofoats (Eppendorfer, 1978; Saastamoinen, 1987).

Oil content

Several factors affected the oil content of the testcultivars Puhti and Veli (Table 6). The main factoraffecting oil content was the average temperature ofthe growing period. A low average temperature in-creased the oil content of oats. Significant positivecorrelations were found between oil content and pre-cipitation (Table 3). Precipitation was not, however,significant in explaining oil content. The effect of lowtemperature in increasing the oil content of oats hasalso been described previously (Beringer, 1967;Welch, 1975; Saastamoinen et al., 1989, 1990;BJ0mstad et al., 1994). Soil nutrient levels also influ-enced oil content. The oil content of the Puhti varietywas increased by the high soil K level and decreasedby the high Ρ and Mg levels of the soil. In the case ofthe Veli cultivar, oil content was dependent only onthe average temperature of the growing period andthe soil Ρ level.

Hectolitre weight, grain size and hull content

Hectolitre weight of both test cultivars was dependenton precipitation (Table 7). High precipitation de-creased hectolitre weight. The hectolitre weight of theVeli cultivar was also dependent on the soil Mg level.Correlation coefficients between hectolitre weight andaverage temperature of the growing period were sig-

Table 5. Dependence of protein content (y) on precipitation (x) in Veli and Puhti varieties in the years 1987-90analysed by multiple step-wise regression analysis

Variety

VeliPuhti

Number

4747

of trials Equation

y= 17.87 -0.016XK= 16.73-0.014x

rMOO

67.6270.20

(%) F-Value for

37.90***43.72***

model

f-Value for:

X

-6.16***-6.61***

Constant

30.58***32.21***

***P<0.001.

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Grain yield and quality of oats

nificantly positive (Table 3), but the average tempera-ture of growing period was not a significant factor inexplaining the variation in hectolitre weight (Table 7).

Ρ fertilization was the most significant factor ex-plaining the variation in the 1000 grain weight of thetest varieties (Table 8). High Ρ fertilization decreasedthe 1000 grain weight of both varieties. High precipi-tation was a decreasing factor in the 1000 grain weightfor Puhti, but not for Veli. High soil pH increased theseed size of Veli. Hellewell et al. (1996) found a 51%higher grain weight with 15°C day/15°C night thanwith 31°C day/15°C night temperature in a growthchamber experiment. The average temperature of thegrowing period was not, however, significant in ex-plaining the variation in grain size found in the presentstudy.

The effect of Ρ fertilization in decreasing oat seedsize can perhaps be understood by the effect offertilization in increasing the floret number of the oatpanicle. The number of seeds increases but seed sizesimultaneously decreases. Furthermore, it has beenfound that in Finland the 1000 grain weight of oats ishigher in organic culture than by normal cultivationwith mineral fertilization (unpublished results). In oatsa lower nutrient level appears to increase the seed size.

Environmental factors had different effects on thehull contents of the test cultivars (Tables 3 and 9). Ahigh mean temperature of the growing period andhigh Ρ fertilization increased the hull content of Velisignificantly. The hull content of Puhti was dependenton the soil Mg level. A high temperature decreased theseed size and increased the hull content of the Velivariety. Ρ fertilization had the same effect, in decreas-ing seed size and increasing hull percentage.

Conclusions

The results suggest that the grain yield of oats is notincreased by a higher temperature during the growingperiod. In addition, the reaction of the test varieties tothe temperature during the growing period varied. Theearly variety Veli is adapted to grow in low tempera-tures. This variety produced higher yields at lowertemperatures during the growing period. Yields ofPuhti, a mid-early variety, were increased by high Ρfertilization. The possibly elevated atmospheric CO 2

concentration and higher average temperature of thegrowing period in the future would create a need foroat varieties better adapted to the changing conditionsin northern cultivation areas. However, according tothe results of this research and previous reports onoat, oat does not appear to adapt well to cultivationin a warmer climate.

The oil content of both test varieties, Puhti andVeli, was increased by low temperatures during the

135

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Table 7. Dependence of hectolitre weight (y) on precipitation (χ,) and Mg level of the soil (x2) in the Puhti andVeli varieties

Variety

VeliPuhti

Numberof trials

4747

Equation

y = 61.51 -0.039x1-0.0078x2

y = 57.87-0.042^

Λ2 · 100

47.6937.42

(%)F-Value formodel

20.05***26.91***

f-Va lue for:

-5.65***-5.19***

x2

-3.55***

Constant

35.50***29.33***

***P<0.001.

Table 8. Dependence of 1000 grain weight (y) on Ρ fertilization (χΊ), precipitation (x2) and pH of the soil (x3) inPuhti and Veli varieties in a 4-year study

Variety

VeliPuhti

Numberof trials

4747

Equation

K=31.19-0.y= 43.19-0.

16x, + 1.41x3

18x,-0.015X2

Γ2·100

20.2228.06

(%)F-Value forequation

5.58**8.58***

f-Value

-3.02**-2.66*

for:

x2

-2.55*

x3

2.10*

Constant

8.16***17.89***

* P<0.05; ** P<0.01; *** P<0.001.

Table 9. Dependence of hull content (y) on Ρ fertilization (x,), mean temperature of growing period (x2) and Mg

level of the soil (x3) in Veli and Puhti varieties in a 4-year study

Variety

VeliPuhti

Numberof trials

4747

Equation

K=9.56 + 0.18x1 + 0.62x2

y =20.86 + 0.0080x3

r2-

53.41.

100

6734

(%)F-Value forequation

8.70***9.28**

f-Value

3.39**

for:

x2

2.85**3.05**

Constant

2.54*20.11***

* P<0.05; ** P<0.01; *** P-cO.001.

growing period. The protein content of both cultivarswas decreased by high precipitation. The mean tem-perature of the growing period was not significant inexplaining the variation in protein content, even if thecorrelation coefficients between the protein contentand mean temperature of the growing period weresignificantly positive.

The effects of environmental factors on grain size,hull content and hectolitre weight were more compli-cated. High precipitation was the main factor causingdecreased hectolitre weight in both varieties. Grainsize was mainly decreased by high Ρ fertilization inthe case of both varieties. Hull content was decreasedby high temperature and Ρ fertilization in the Velivariety and by a high level of Mg in the soil in thePuhti variety.

Acknowledgements

This research was carried out at the Institute of PlantBreeding, Agricultural Research Centre of Finland. Itwas financed by the Finnish Ministry of Agriculture

and Forestry, whose support is gratefullyacknowledged.

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