Growth and nutrient uptake of coniferous seedlings: Comparison among 10 species at various seedbed densities

Plant and Soil 38, 125-143 (1973) Ms. 1979

GROWTH AND NUTRIENT UPTAKE OF CONIFEROUS SEEDLINGS : COMPARISON

AMONG 10 SPECIES AT VARIOUS SEEDBED DENSITIES *

by NORMAN A. RICHARDS, ALBERT L. LEAF,

and DONALD H. BICKELHAUPT **

SUMMARY

The evaluation of biomass production and uptake of N, P, K, Ca, and Mg for various plant components (roots, stems, and foliage) and totals by 10 species of 2-O coniferous seedlings grown at a controlled range of densities in a highly productive forest nursery documents considerable differences among species and seedling parameters. The species are ranked by biomass and nutrient-element relationships on a unit area of seedbed basis, quanti- fying the magnitude of the differences among the species at the various density levels. The 10 species include Abies balsamea, Larix leptole+Gs, Picea abies, Picea glauca, Picea mariana, Picea pungens, Pinus resinosa, Pinus strobus, Pinus sylvestris, and Pseudotsuga menziesii.

INTRODUCTION

Differential nutrient element uptake is known to exist among plant species growing on a given site. These differences in uptake have been attributed to genetic characteristics of the plants, and to the total physical, chemical, and biological aspects of the environment to which the plants are subjected. Research involving tree species - nutrient element relations in this regard has been relatively minimal as evident in reviews by D a v e y 1 and V o i g t 8. Though selected tree species grown under controlled environment

* Contribution of the Forest Resources Council, School of Environmental and Re- source Management, State University of New York College of Environmental Science and Forestry, Syracuse, New York 13210.

** Research Associate in Silviculture, Professor of Forest Soil Science, and Research Assistant in Silviculture, respectively.

126 N. A. RICHARDS, A. L. LEAF AND D. H. BICKELHAUPT

conditions have received intensive study 4 5, and a general qualitat- ive rating of species as to their site requirements is known, there is little comparative data among species for such ratings 2 3 9.

The purpose of this paper is to quantify growth and nutrient element uptake of seedlings of several coniferous tree species over a controlled range of seedbed densities. At the seedling stage in an intensively managed nursery, the three species are contained in a relatively homogeneous physical environment and, thus, can be subjected to a more simplified evaluation and interpretation than forest stands with extensive rooting patterns established within the heterogeneous soil medium.

METHODS AND MATERIALS

At the Syracuse Forest Experiment Station nursery, following a summer harvest of a crop of 3-O (Picea glauca (Moench.) Voss., the 0 to 15 cm soil depth conditions ranged as follows: Texture: 83-87% sand, 7-9% silt, 5-7% clay; Organic matter: 4.7-6.0%; pH value: 5.7-6.9; Total N (Kjeldahl proce- dure) : 0.115-O. 165% ; Available P (0.03N NHdF in 0.025N HCI extractable) : 16-26 ppm; Exchangeable K (1N NH4 OAc, pH 6.8 extractable) : 35-64 ppm; Exchangeable Ca (1N NH4 OAc, pH 6.8 extractable) : 1480-1830 ppm; Exchangeable Mg (1N NH4 OAc, pH 6.8 extractable) : 76-96 ppm.

In autumn preceeding establishment of the current study, the seedbed area received 224 kg/ha of ammonium sulfate, 560 kg/ha powdered sulfur, and a top dressing of peat moss at 630 cu m/ha. This was all tilled into the soil be- fore seeding the 10 species of conifers at various densities. Just prior to the beginning of the second year of growth of the seedlings of this study, a top dressing of 5-10-5 commercial fertilizer was applied at the rate of 1 .O kg/l0 sq. m of seedbed. Irrigation water supplemented precipitation throughout this study so the soil received at least 25 mm of water per week during the growing seasons. These conditions represent a favorable nursery soil condition for the 10 coniferous tree species studied: Abies balsavnea (L.) Mill.; La+& leptolepis (Sieb. and Zucc.) Gord.; Picea abies (L.) Karst.; Picea glauca (Moench.) Voss.; Picea rnaviana (Mill.) B.S.P.; Picea pungens Engelm.; Pinus resinosa Ait. ; Pinus stvobus L. ; Pinus sylvestris L. ; Pseudotsuga menziesii (Mirb.) France.

Each species of known seed source was sown at several densities per unit area, generally six densities ranging from a low of 100 to 250 seedlings/sq.m to a high of 700 to over 1500 seedlings/sq.m of seedbed. Growth of species was typical of that experienced in this nursery except for Picea abies which has been one of the faster growing Picea species.

In the autumn, after two years from seeding, 50 to 100 seedlings, depending on their size, were carefully lifted in toto from each species and density com- bination. These seedlings were separated into foliage, stem (all wood anp

GROWTH AND NUTRIENT UPTAKE OF SEEDLINGS 127

bark above-ground plant components including branches if present), and root components, measured for 1-O and 2-O height, 2-O root collar diameter, dried to constant weight at 70°C weighed, ground to pass a 2-mm sieve, and analyzed for ash, N, P, K, Ca, and Mg by plant components. Nitrogen was determined by the macro-Kjeldahl technique, and the other elements were analyzed from samples dry-ashed at 480°C for 15 hours and taken up in weak HCl solution. Phosphorus was determined by the molybdo-vanadate proce- dure with a spectrophotometer, K was determined with a flame spectrophotometer, and Ca and Mg were determined with an atomic-absorption spectrophotometer.

Since the range of seedbed densities in the nursery was not identical for each species, the data that foliows are based on values calculated from simple regression equations. The form of the regression equation was delimited by the 0.10 level of significance to obtain the best fit of the data. Therefore, all calculations were first done using

Y = a + blogX,

and, if the ~2 values denoted non-significance, the calculations were repeated using

Y=a+bX,

where X was seedbed density in numbers/sq.m, and Y was each of 24 individual characteristics of each species, e.g. total height; second year height; root collar diameter; dry weight of roots, stems, and foliage; ash, N, P, K, Ca, and Mg contents of roots, stems, and foliage. Of the 240 regressions, it was necessary to use the linear regression in only two cases, and in six cases where regression equations were nonsignificant, mean values were used at all densities.

INDIVIDUAL SEEDLING DATA

Results of this study may be expressed either on a per seedling or on a per unit area of seedbed basis. The first method indicates the reduction in individual seedling dimensions and nutrient element uptake with increasing seedbed density while the latter expression indicates the increased dry matter production and nutrient element uptake with increasing seedbed density. There is an inverse relation between individual seedling development and yields per sq. m of seedbed in relation to seedbed density, with considerable differences among species in the specifics of this relationship.

Ideally, over a full range of seedbed densities, the curve of individual seedling growth and nutrient uptake would demonstrate three zones as schematically shown in Figure 1. Zone A denotes a low density having little effect on individual seedling yield. Zone B de-


INDIVIDUAL SEEDLING DRY MATTER PRODUCTION, AND NUTRIENT UPTAKE

A

B

\ ? I

I

L I

- SEEDBED DENSITY - Fig. 1. Schematic diagram of individual seedling development in relation to

seedbed density. Zones A, B, Cl and C2 explained in text.

notes the portion of the curve where increasing density results in a sharp decrease in individual seedling yield due largely to decreased seedling branching. At the inflection between zones B and C an un- branched seedling results. Zone C denotes the portion of the curve where increasing density results in a gradual decrease in seedling size and nutrient element uptake due largely to reduced diameter growth in Cl and also reduced height in C2, as well as decreased root development.

Not all 10 coniferous species in this study show all zones depicted in Figure 1 because of the range of densities used and growth characteristics of the species. Abies balsamea and Pseudotsuga mevziesii show zone C only because these two species normally are not branched as 2-O stock. Lark lefitolepis shows zone C only because at 20 seedlings/sq m zones A and B have already been surpassed. In all other species, except Picea mariana, zones B and C were evident, but zone A had been surpassed at 200 seedlings per sq.m. Only Picea Ynariarza showed all three zones. In general, the inflection point between zones B and C occurred below 400 seedlings per sq.m. for Pinus species and above this density for Picea species. Picea mariana showed zone A ending at approximately 400 seedlings per sq.m.


TABLE 1

Mean 2-O seedling diameter, height and dry weight of 10 coniferous species grown at various seedbed densities.*

2-O diameter at 2-O total height 2-O total dry root collar, mm growth, cm weightlsdlg., g

Density, no./sq.m 300 600 900 1200 300 600 9001200 300 600 900 1200

Species Larix leptolepis Pinus sylvestris Pseudotsuga rnenziesii Pinus strobus Pinus resinosa Picea pwgens Picea nzariana Picea glauca Picea abies Abies balsamea

4.3 3.9 3.8 [3.7] 71 65 61 [58] 4.2 3.6 3.0 [2.6] 4.1 3.4 [3.0 2.71 27 24 [22 211 3.6 2.5 Lt.9 1.61 2.8 2.7 2.6 2.5 37 37 37 37 1.8 1.9 1.7 1.5 2.9 2.5 2.2 [2.0] 29 29 29 [29] 2.3 1.7 1.3 [l.l] 2.8 2.5 2.3 [2.2] 31 22 20 [18] 1.9 1.5 1.2 [l.O] 3.3 2.8 2.4 2.2 38 36 35 34 1.6 1.4 1.2 1.1 3.0 2.4 2.2 2.0 40 38 37 36 2.2 1.4 1.1 0.9 3.2 2.6 2.3 2.1 35 34 33 32 1.5 1.2 1.0 0.9 2.8 2.3 2.0 1.9 33 33 33 33 1.5 1.1 0.9 0.8 2.7 2.4 2.2 2.1 31 28 27 26 0.5 0.6 0.6 0.5

* Data in brackets are extrapolated by regression analysis beyond actual densities at which seedlings were grown in the nursery.

The ranges in diameter at root collar, total height and average seedling dry weight for 2-O stock at various densities are shown in Table 1. Generally, the decrease in individual seedling yield and nutrient uptake with increased seedbed density are not well depicted in seedling height and diameter measures because of the effect of density on seedling branching. Except in the case of Lark leptolepis, which outgrew all other species approximately three-fold in height the first year from seeding, there were no significant relationships between first year height growth and seedbed density among species tested. There was, however, evidence of relationships between 2-O total height, second year height growth, 2-O collar diameter, and seedbed density as noted for most of the species.

UNIT AREA OF SEEDBED DATA

Biomass relationships

As is apparent from Figures 2 through 5 there were striking differences among species at various densities in total biomass production and proportions in foliage, stem and root components. At a density in excess of 1000 seedlings per sq.m, a five-fold difference in total


LARIX LEPTOLEPIS

PINUS SYLVESTRIS

PSUEDOTSUGA MENZIESII

PINUS STROBUS

PINUS RESINOSA

PICEA PUNGENS

PICEA MARIANA

PICEA GLAUCA

ABIES BALSAMEA

% b b ;: tT g -- - -z --

000000 Q s

:: 0

SEEDUNGS PER SQ. M.

Fig. 2. Root dry-weight production per unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.

biomass existed between the largest values, La& leptolepis with almost 30 metric tons per ha, and the least values, Abies balsavnea with approximately 6 metric tons per ha. At a density of 300 seedlings per sq. m, the range in total biomass for these two species was greater than 12-fold, but the actual dry matter production was considerably lower than at the highest density. The data for all other species investigated were between these extremes.

From low to high density, total biomass for each species increased from less than two-fold, e.g. P&m sylvestris, Pinus strobus, Pinm resinosa, and Picea mariana, to over three-fold for Pseudotswga men- xiesii and Abies balsamea.

Of this total biomass, less than 30 per cent was in the root component for the Pinus species, Lark lefitolepis, and Picea mariana, while up to 35 per cent was in this component for the other Picea spe-


PSI

LARIX LEPTOLE

JEDOTSUGA MENZIESI I

PINUS STROBUS

PINUS RESINOSA

PICEA PUNGENS

PICEA GLAUCA

PICEA ABIES

ABIES BALSAMEA

SEEDLINGS PER SQ. M.

Fig. 3. Stem dry weight production per unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.

ties and Pseudotsuga nzenziesii. Abies balsamea had thehighest proportion, approximately 40 per cent, of total biomass in roots. Increasing density of seedlings did not greatly change proportions of total biomass in roots. This proportion tended to remain relatively constant or slightly decrease, e.g. Picea glauca, with increasing density.

Of the total biomass, the stem component ranged from less than 20 per cent for Piws resinosa and Pirzus strobus to almost 60 per cent for Larix leptolepis. Except for the Lavix species the stem data was less than 40 per cent of the total biomass. Increasing density of


PSUE

LARIX LEPTOLEPIS

PINUS SYLVESTRIS

:DOTSUGA MENZIESI I

PINUS STROBUS

PINUS RESINOSA

PICEA PUNGENS

PICEA MARIANA

PICEA GLAUCA

PICEA ABIES

ABIES BALSAMEA


Fig. 4. Foliage dry weight production per unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.

seedlings had little effect in changing proportions of total biomass in stems, except for a proportional increase for Picea glauca.

With the exceptions of Abies balsavnea and Larix leptolepis, the greatest proportion of total biomass was in the foliage component. The highest proportion in the foliage component was in the Pinus species, approximately 50 to 65 per cent followed by 40 to 50 per cent in the Picea species, with Psetidotsuga Ynenziesii averaging 35 per cent, Abies balsamea averaging 30 per cent, and Lark leptolepis averaging 25 per cent of the total biomass. Changes in the foliage component proportion of total biomass with increasing density was relatively minimal except for porportional increases for Abies balsamea and Picea pungem, and a proportional decrease for Pintis resinosa.

PSI


- ”

LARIX LEPTOLEPIS

PINUS SYLi’ESTRlS t

,,,$jc ;+

JEDOTSUGA MENZIESII

PINUS STROBUS

PINUS RESINOSA

PICEA PUNGENS

FICEA MARlANA

PICEA GLAUCA

PICEA ABIES

ARIES BALSAMEA


Fig. 5. Total dry weight production per unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.

Nutrient element relationships

As evident in Tables 2 through 5, the general patterns of nutrient element uptake by various species, plant components, and densities were generally related to biomass production.

The largest biomass production with Lark leptolepis at over 1000 seedlings per sq. m, approximately 30 metric tons/ha, contained almost 16 metric tons/ha of ash, 350 kg per ha of N, 90 kg/ha of P, 220 kg of K per ha, 160 kg of Ca per ha, and 35 kg of Mg per ha. Con- versely, the lowest total biomass production with Abies balsamea at 300 seedlings/sq m, contained approximately 180 kg of ash per ha, 24 kg of N per ha, 3 kg of P per ha, 8 kg of K per ha, 4 kg of Ca per ha, and 1 kg of Mg per ha.


TABLE 2

Total and plant component dry weight and content of ash/unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.*

Dry weight, kg/ha ilsh content, kg/ha

Density, no./sq. m 300 600 900 1200 300 600 900 1200

La+ leptolepis Roots 2284 Stems 7437 Foliage 2790 c 12512

Pinus sylvestris Roots 1592 Stems 3576 Foliage 5568 x 10735

Pseudotsztpa nzetiziesii

126 168 183 317 225 500 534 984

192 395 655

1242

88 107 141 208 232 338 - - 461 652

119 247

I-

399 765

127 274 443 - a44

3509 4674 3954 5311 3796 4986 __ -

11259 14971

Roots _ 1518 Stems 1633 Foliage 1761

c 4912

Pinus strobus Roots Stems Foliage c

1890 1145

3768 6804

Pinus resinosa Roots Stems Foliage

c

1109 1955 935 1585

3808 5291 5852 8830

Picea pulzgens Roots Stems Foliage c

1242 2085 1535 2819 1649 3630

4426 a533

Picea mariana Roots Stems Foliage I:

1148 2212 2834 6194

Picea glauca Roots Stems Foliage c

1532 1966 720 i 780

2195 3737 4447 7483

Picea abies Roots stems Foliage

c

1546 2388 2880 3229 1172 I 728 2052 2283 1749 2754 3341 3753

4467 6869 8274 9270

Abies balsanzea Roots Stems Foliage c

503 400 100

1003

2864 3437 1654 1951 5490 6497 __~

10010 11886

2450 1965 2234 6158

10572

-1 2801

6774 ,11809

5500 33 6274 54 5831 102

17606 188

2577 2927 3570 4102 4789 5611 - ~

10936 12641

1442 1615 1737 3034 3514 3855 3851 4446 4868 -__- 8327 9575 10460

2220 240 1 2430 2891 4639 5279 -~ 9289 10571

1511 2101 2320 1108 1522 1816 1015 1550 1930 ____- 3634 5174 6266

120 129 228 - 477

127 46

172

345

202 66

251 - 520

47 81 42 65

153 218 241 364

ii 96 -

193

78

2% - 416

41 62 67 107

202 258 310 426

60 45

192 296

85

2;; - 445

84 40

144 - 267

170 55

193 83

- -

48 47 a5 -

180 -

171 207 173 204 302 354 646 zz

246 78

297 621

101 79

256 - 436

115 89

283 m I

92 131 323 546

103 154

382 - 639

73 82 130 147 291 314 - - 494 542

99 110 104 120 328 364 - - 532 594

221 257 64 71

222 242 - - 507 570

77 63

133 273

97 74

167 - 338



Ash content. From low to high density, total ash content increased from slightly less than two-fold to approximately four-fold. The magnitude of this range by species is generally similar to the biomass results. However, ash content increased with greater density at a more rapid rate than biomass in foliage of Lark leptolepis, in roots and total content of Pseudotsuga Yuzenziesii, in stems, foliage and total content of Picea @figens, in roots and stems of Picea maviana, and in roots of Picea a&es. Conversely, ash content increased with greater density at a less rapid rate than biomass in roots of Lark leptolepis and Picea pungens, in foliage and total content of Picea glauca, and in foliage of Picea abies.

The proportions of total ash in the root component ranged from approximately 15 to 20 per cent of the total ash for Pintis sylvestris, Picea mariana, Picea glnz~ca, and Lavix leptolepis, to a high of 35 to 45 per cent for Picea abies. The proportions of total ash in the stem ranged from a low of 10 to 15 per cent for Picea abies and Piws stvobus to a high of approximately 30 to 35 per cent for Larix leptolelpis and Pinus sylvestris. As would be expected, foliage contained the highest proportion of ash of the seedling components, ranging from 40 to over 60 per cent of the total ash. Picea glauca and Picea mariana contained the highest proportion of foliage ash, approximately 60 to 65 per cent, while Pinus stvobus, Pseudotsuga menziesii, and Abies balsamea ranged from 40 to 50 per cent. Re- lative to biomass, all species except the three Pinus species had a higher proportion of total ash in foliage, and only Pings strobes had a lower proportion in foliage, but higher proportion in roots. The Lavix species and three of the Picea species had lower proportions of total ash relative to biomass in stems, while Abies basamea and Pseudotsuga menziesii had lower proportions in roots.

N content. From low to high density, total N content increased from extremes of 1 &fold for Picea abies to over three-fold for A bies balsamea. Nitrogen content increased with greater density at a more rapid rate than biomass in foliage of Larix leptolepis, Pseudotsuga menziesii and PinGs resinosa, and in stems of Pinus sylvestvis and Picea @ngens. Conversely, N content increased with greater density at a less rapid rate than biomass in roots of Pinus Yesinosa, in roots, foliage, and total content of Picea pungens, in roots, stems, and total content of Pseudotsuga menziesii, Picea abies and Abies balsamea.


TABLE 3

Total and plant component content of N and P/unit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.*

N content, kg/ha P content, kg/ha

Density, no./sq.m 300 600 900 1200 300 600 900 1200

Larix 1epto1cpis Roots 21 Stems 102 Foliage 21 r, 144

Pinus sylvestris Roots 11 Stems 34 Foliage 102 c 147

PsezGdotsuga menziesii Roots Stems Foliage

Lx

11 18 26 57;

Pinus strobus Roots Stems Foliage

z

Pinus resinosa Roots Stems Foliage r,

Picea pungens Roots Stems Foliage c

Picea mariana Roots Stems Foliage c


Piceaabies Roots Stems Foliage ;r:

Abies balsamea Roots Stems Foliage B

22 29 36 46 62 83 - -

120 158

17 16 73 -

106

:3’ 105 -

155

9

4; 63

13 16 16 20 69 83 55 120

9

374 50

13

:96 88

11 34 48 94

13 9

28 5T

7 14 9 16 7 22

iz 33

5% - 251

15 56

130 201

4.5 189

80 - 314

1% 70

146 234

32

12 - 184

16 26 69 -

111

13

2 - 109

15 26 67 -

109

18

3”; 75

34 53 98

185

36 31

0

137 204

17 32 78

127

14 45 60

119

24 12 42 78

4 21 18

43

3

1; 24

: 8

14

t 10 18

2 2 7

lo

3” 6

ii

42 9

is-

2

1; 16

2

6” ii

1

t 3

28 67

10 36 34 so

5 13 16

34 i

5 16 19

40

6 18 20 44

3 9

14 -5%

5

1: 26

‘: 18 Fl

3

142 19

4

12 24

2

1;’ 18

z 14

22

2 6

16 25

2 3 3 6 6 7

12 14 15 20 23 25

3

12 24

3 7

19 T9

: 11 .zT

: 5

E

4 a

21 32

7

1: 23

6” 6

16

* Data in brackets a re extrapolated by regression analysis beyond actual densities at which seedlings were grown i n the nursery.


Of the total N content, the proportion in the root component ranged from approximately 10 to 30 per cent for Pinus sylvestris and Abies balsamea, respectively. The proportions of total N in stems ranged from 15 per cent for Pinus strobes to approximately 60 to 70 per cent for Lark leptolepis. However, with the exception of the Lark species the stem N was less than 40 per cent of total N content. With the exceptions of Lark leptolepis and Abies balsamea, all species contained over 50 per cent of their total N in foliage. Relative to biomass, all species, except the Larix, had a higher proportion of total N in the foliage, whereas the Lark had a higher proportion in stems. All species had a lower proportion of total N relative to biomass in roots.

P cant ent. From low to high density, total P content increased from slightly less than two-fold for Pinus sylvestris, Pinus strobus, and Picea mariana to approximately five-fold for Abies balsamea. Phosphorus content increased with greater density at a more rapid rate than biomass in all components of Pinzts resinosa, in roots of Picea abies, and in stems of Abies balsamea. Conversely, P content increased with greater density at a less rapid rate than biomass in total content, stems and foliage of Lark leptolepis, Pseudotsuga menziesii, and Picea glauca, in all components of Picea pungens, in stems of Picea abies, and in total content, roots, and foliage of A bies balsamea.

Of the total P content, the proportion in the root component ranged from 10 to 30 per cent, with most of the species averaging less than 20 per cent of the total P in roots. The proportions of total P in stems ranged from slightly less than 20 per cent for Picea abies to a high of 45 to 50 per cent for Lark leptolepis. With the exceptions of the Lark species, Abies balsamea and Pinus sylvestris, all species contained over 50 per cent of their total P in foliage. Relative to biomass, all species, except the three Pinus species, had a higher proportion of total P in foliage; and all, except Pinus sylvestris, had a lower proportion in roots.

K content. From low to high density, total K content increased from approximately l&fold for Picea wtariana and Picea abies to over four-fold for Abies balsamea. Potassium content increased with greater density at a more rapid rate than biomass in foliage of


TABLE 4

Total and plant component contents of K, Ca, and M&nit area of 10 species of 2-O coniferous seedlings grown at various seedbed densities.*

K content, kg/ha Ca content, kg/ha Mg content, kg/ha

Density, no./sq. m

Larix le~tole~is Roots Stems Foliage

Iz

Pinus sylvestris Roots SkIlS

Foliage c

Pseudotsuga menziesii Roots Stems Foliage

c

Pinus strobus Roots Stems Foliage c

Pinus resinosa Roots Stems Foliage r,

Picea pungens Roots Stems Foliage

c

Picea mariana Roots Stems Foliage r,


Picea abies Roots Stems Foliage c

Abies balsa,hea Roots Stems Foliage c

300 600 900 1200 -

300 600 900 1200 300 600 9001200

15 45 24 84

10 29 44

33

2 10 18

30

9 22 29 34 8 17 21 25

20 37 47 54 t 75 98

- 113

1: 28

r

24; 364 40 7

34 44 51 64 s4 99

1: 18 7 2: 41

-Ii

49 55

63 75 94

5 7 a

49 70 83

3 6

26 36

4 5 5 6 6 13 16 19

1: ii 2 $: 2: i: :: 21 61

28 54 ?6 81 32 67 -- 87 101

2 12 27 41

6 9 10 11

3; 48 11 54 12 L”9 51 67 77 83

2

149 26

9 12 14 15 14 19 22 24 30 54 68 78 - --- 53 a5 103 116

2 7

16 26

4 130 1’: 12 22 25 28 35 40 44

11 19 23 27 2 3 6 9 11 12

f 2

23 34 40 44 3 4 5” 40 62 ;i;l 83 s 8 9

3 4 1; 6 3 9 2 9 13 t: 8 zz 31 37

1 2 3 4 2 4 5 6 3 6 8 8 6 12 16 18

2 3 4 2 2 3 6 8 9

10 14 16

1 2 2 2 3

: 8 9 --- 7 12 15

; 2

3” A 4 5 b

a -- 9 11

1 1 2

: 2 6 3 6 ?

-- 9 10

1 2 2 2 3 4 4 6 7 7 ii 13

Tr. 1 2 Tr. 1 2 Tr. 2 2

146

‘4 3

1 $1



Larix leptolepis and Picea jwmgens, in roots and stems of Pseudotsuga menziesii, in roots of Pings resinosa, and in sterns of Abies balsauuzea. Conversely, K content increased with greater density at a less rapid rate than biomass in roots of Lavix le@ole$s, in foliage of Pseudot- saga ynenziesii, in roots and stems of Picea #wagens and Picea glauca, in all components of Picea abies, and in total content, roots and foliage of Abies balsamea.

Of the total K content, theproportionin the root component wasless then 1 Oper cent forPsetidotsugavnenziesii and the Picea species, and 10 to 25 per cent for the remaining species. The proportions of total K in stems ranged from slightly less than 20 per cent for Pinus resinosa to a high of 45 to over 55 per cent for Lavix leptolepis. With the exceptions of the Lark and Abies balsamea, all species contained over 50 per cent of their total K in foliage. Pinus resinosa averaged approximately 70 per cent of its total K content in foliage. Relative to biomass, all species, except Pinus sylvestvis, had a higher proportion of total K in foliage, and all species had a lower proportion in roots.

Ca content. From low to high density, total Ca content increased less than two-fold for Pinus sylvestvis, Pinus styobus, and Picea Ynaviana, to approximately threefold for Pseudotsuga menziesii and Picea flungens. Because of the very low value at low density, total Ca content of Abies balsamea increased considerably in the same density range. Calcium content increased with greater density at a more rapid rate than biomass in total content, roots and stems of Lark leptolepis and Picea pungen, in all components of Pinus resinosa and Abies balsawea, and in foliage of Picea rnariana. Con- versely, Ca content increased with greater density at a less rapid rate than biomass in total content and foliage of Pious sylvestris, in total content, stems and foliage of Pseudotsuga menziesii, in foliage of Picea pungens, and in stems of Picea glauca.

Of the total Ca content, the proportion in the root component ranged from 10 to 30 per cent, with half of the species averaging less than 20 per cent. The proportions of the total Ca in stems also ranged from 10 to 30 per cent, with half of the species averaging less than 25 per cent. All species contained over 50 per cent of their total Ca contents in foliage, with Picea mariana ranging from 70 to 75 per cent in this component. Relative to biomass, all species had a higher proportion of total Ca in foliage, and, except Picea glauca, had a


lower proportion in stems. All species, except the three Pinus species, had a lower proportion of total Ca relative to biomass in roots.

Mg content. From low to high density, total Mg content increased from l&-fold for Picea mariana to three-fold for Pseudotsuga ruzenziesii. Because of the very low value at low density, total Mg content of Abies balsamea increased considerably in the same density range. Magnesium content increased with greater density at a more rapid rate than biomass in roots of Lark leptolefiis, Pseudotswga menziesii and Picea abies, in roots and foliage of Pinus sylvestris, in roots and stems of Piws resinosa, and in total content, roots and stems of Abies balsamea. Conversely, Mg content increased with greater density at a less rapid rate than biomass in foliage of Larix leptolepis and Abies balsamea, in stems of Pinus sylvestris and Picea mariana, in total content, stems and foliage of Pseudotsuga menziesii, in total content, roots and foliage of Picea pungens, in total content and stems of Picea glauca, and in stems and foliage of Picea abies.

Of the total Mg content, the proportion in the root component ranged from less than 15 to over 30 per cent, with over half of the species averaging less than 20 per cent. The proportions of the total Mg in stems ranged from over 15 to approximately 50 per cent, with over half of the species averaging approximately 30 per cent. With the exceptions of Larix leptolepis and Abies balsamea, all species contained over 50 per cent of their total Mg contents in foliage. Relative to biomass, all species, except Pinus sylvestris, had a higher proportion of total Mg in foliage, and a lower proportion in roots.

CONCLUSIONS

The evaluation of biomass production and nutrient element uptake by 10 species of 2-O coniferous seedlings grown at various densities in a highly productive forest nursery documents considerable differences among species. Generalizing over the range of densities, the ranking of species in decending order of total production indicates:

Lavix leptolepis > Pinus sylvestris > Pseudotsuga menziesii > Pilzus strobus >I= Pinus resinosa = Picea pungens = Picea mariana = Picea glauca >I= Picea abies > Abies balsamea.


TABLE 5

Ratios of total and plant component dry weight and contents of ash, N, P, K, Ca and Mg/ unit area of 10 species of 2-O coniferous seedlings at 1200 plants/sq.m to 300 p1antslsq.m

Species and component

Dry Ash N P K Ca Mg weight content content content content content content

- Larix leptolepis

Roots Stems Foliage z:

2.7 1.7 2.3 2.5 2.8 3.4 2.5 2.7

Piws sylvcstris Roots Stems Foliage Ix

Pseudotsuga menziesii Roots Stems Foliage c

Pitius strobus Roots Stems Foliage c

Pilzus resinosa Roots Stems Foliage 2

Picea pungens Roots Stems Foliage c

Picea marialza Roots Stems Foliage c

Picea glazlca Roots Stems

1.5 1.4 1.7 1.8 1.5 1.5 1.8 2.0 2.0 2.3 2.1 2.1 1.9 1.6 1.7 1.9 1.6 1.8 1.7 1.1 2.1 1.8 1.8 1.7 1.9 1.8 1.3 1.8

3.6 6.3 3.8 3.8 3.3 3.5 3.6 4.1

2.0 1.9

t :;

2.5 2.5 2.4 2.1 1.8 1.9 2.0 2.0

2.4

32.47 2:8

Foliage B

Picea abies Roots Stems Foliage c

Abies balsamea Roots Stems Foliage c

2.2 1.9 1.9 2.0

1.9 3.5

34::

2.0 2.2 1.6 1.7

1.8 2.7 1.9

2.0

3.1 1.8 1.7 2.1

-

2.5

:A 2:5

2.8 2.2 1.9 2.2

2.1 3.7 2.1 2.6

3.2

32:; 3.3

3.8 4.6 3.8 2.7 4.0 2.9 2.5 2.8 2.8 2.7 3.4 3.1

2.1 1.9 1.9 1.9

1.9 2.0 1.9 1.9

it 2:1 2.1

3.4

22.68 217

1.9 1.1

4.3” 2:5

22.63 2:3

t :;

f :L

1.6 1.7 1.6 1.6

z 2:2 1.9

t ::

1:;

2.8 1.5

2.0 2.1

2.9 2.5

35::

3.1

2: 5:1

i.7 216 2.8

3.1 2.2 2.2 2.3

E 2:5 3.1

if 2:o 2.0

2.2 1.8

f::

2.0

t78 1:8

2.9 2.3 1.9 2.1

2.2 2:2 2.5

3.3 3.1 1.8 2.2

1.3 3.0 1.8 2.3 4.7 2.7 3.7 2.8 1.8 2.9 3.2 2.0

1.6 1.5

t ::

2.0 1.7 1.6 1.4 1.6 . 1.5 1.6 1.5

1.8 1.6 1.8 2.3 1.7 2.0 2.4 2.6 2.2 2.3 2.2 2.1

1.6 1.7

t :7”

22 614 4.7

2.5 2.0 1.9 2.1

10.2 ̂

2.4 1.6 1.9

2.0

7.7 6.2 7.7

63.4 14.5 15.3 10.7


Because of these differences in growth rates, the effects of seedbed density differ among species. In general, there is a l&- to 3&-fold increase in biomass over a four-fold increase in density, except for Abies balsamea, the slowest growing species.

Total uptake and contents of nutrient elements in the various plant components (roots, stems, and foliage) closely parallelled biomass production, but the plant components showed more variation (Table 5). These data do not separate limiting nutrient element levels from luxury consumption levels, nor do they dinstinguish nutrient element requirements of these species. However, they do quantify the ranges of nutrient element uptake under favorable seedbed conditions.

The distribution of the nutrient elements within the plant is documented. Relative to biomass, a higher proportion of the nutrient elements are in foliage and a lower proportion in roots. How- ever, the many differences among species and nutrient elements may be significant.

Biomass production and nutrient element uptake in coniferous nursery seedbeds is substantial. Estimating that, for most of the species, approximately 65 to 75 per cent of their total two-year growth occurred in the second year, the biomass production and nutrient element uptake in that year of growth are comparable’with corresponding data on a per annum basis in the literature for forest stands of more advanced age and maize crops compiled by Ovington and others 6 7.

ACKNOWLEDGEMENTS

Financial support by the State University of New York College of Forestry at Syracuse is gratefully acknowledged. Appreciation is expressed to Professor J. H. Engelken and Messers. L. Carey, N. I. Lamson, and Mrs. M. G. Osborne for assistance in various stages of this study.

Received May 16,1972


LITERATURE

1 Davey, C. B., Biological considerations in fertilizer evaluation. In: Forest Fertiliza- tion Theory and Practice, Tennessee Valley Authority, Muscle Shoals, Ala. pp: 264- 274 (1968).

2 Fornes, R. IS., Berglund, J. V. and Leaf, A. L., A comparison of the growth and nutrition of Picea abies (L.) Karst. and Pinus resinosn Ait. on a K-deficient site subjected to K fertilization. Plant and Soil 33, 345-360 (1970).

3 Heiberg, S. O., and Leaf, A. L., Potassium fertilization of coniferous plantations in New York. Trans. 7th Intern. Congr. Soil Sci. (Madison, Wise.) 3, 3766383 (1960).

4 Ingest ad, T., A definition of optimum nutrient requirements in birch seedlings. I. Physiol. Plantarum 23, 1127-l 138 (1970).

5 Ingestad, T., A definition of optimum nutrient requirements in birch seedlings. II. Physiol. Plantarum 24, 118-125 (1971).

6 Ovington, J. D., Organic production, turnover and mineral cycling in woodlands. Biol. Rev. 40, 295-336 ( 1965).

7 Ovington, J. C., Heitkamp, D., and Lawrence, D. B., Plant biomass and pro- ductivity of prairie, savanna, oakwood, and maize field ecosystems in central Minneso- ta. Ecology 44, 52-63 (1963).

8 Voigt, G. K., Variation in nutrient uptake by trees. IS: Forest Fertilization Theory and Practice, Tennessee Valley Authority, Muscle Shoals, Ala. pp. 20-27 (1968).

9 Wilde, S. A., Forest Soils. Ronald Press, New York (1958).

Documents

Growth and nutrient uptake of coniferous seedlings: Comparison among 10 species at various seedbed densities