Historic, archived document
Do not assume content reflects current scientific knowledge, policies, or practices.
el Ra tote > Veer 4
A9DG
Cry > United State
Department of Agriculture
Forest Service Intermountain Forest and Range Experiment Station
Research Paper INT-281
July 1981
Monoterpenes of Lodgepole Pine Phloem as Related to Mountain Pine Beetles
Walter E. Cole, E. Park Guymon, and Chester E. Jensen
MONOTERPENE MODELS
(38.5 - D) _ 28 0.49
Ra GOO 92 sae
THE AUTHORS
WALTER E. COLE is Project Leader of the Population Dyna- mics of the Mountain Pine Beetle research work unit in Ogden, Utah. This unit was started in 1960 under his direc- tion, as was the early research groundwork on the mountain pine beetle. Prior to this assignment, he did popylation dynamics research, control, and survey work on the spruce budworm and pine butterfly in southern Idaho. He did biolo- gical research and survey data collection on the spruce bark beetle in Fort Collins, Colorado. He began his career with Forest Insect Investigations, Bureau of Entomology and Plant Quarantine, as supervisory control and survey aid in Berkeley, California. Dr. Cole has authored 31 publications.
E. PARK GUYMON is Professor of Chemistry at Weber State College, Ogden, Utah. He received his Ph.D. in analytical and inorganic chemistry from Utah State University. His research activities have covered many interesting and practical aspects in the analysis of natural products and nitrogen fixa- tion. Dr. Guymon’s publications are well-known in these fields of interest.
CHESTER E. JENSEN, retired, served as principal statistician for the Intermountain Forest and Range Experiment Station from 1967 to 1980. He held the same position at the North- eastern and Central States Forest Experiment Stations prior to coming to the Intermountain Station.
RESEARCH SUMMARY
Phloem samples taken from 86 healthy lodgepole pine trees at three points in the 1975 growing season were analyzed for content of dry matter, starch, various forms of sugar and nit- rogen, and of selected monoterpenes. Means for July 10 and 31 were significantly lower than those of June 6 for dry matter, soluble reducing sugars, nitrogen, and monoterpenes. Starches and other sugars were higher. 8-phellandrene was, by far, the most prevalent of the monoterpenes. Dry matter in the phloem contained an extremely small amount of monoterpene by weight but, of this, individual monoterpenes were distributed in about the same proportions found in pure oleoresin by other researchers. Monoterpene contents from the last (July 31) sam- ples were significantly, although weakly, related to the linear positive effects of phloem thickness and radial growth. An inter- active hypothesis is developed for terpene content as a function of phloem thickness, radial growth, and tree diameter. Here, high concentrations of monoterpenes coincide with larger tree diameters, the expected region of high mountain pine beetle survival.
CONTENTS Page INTIRODUGTION 332455. cc eee eee 1 MATERIALS AND METHODS? eee aacee oe eee 1 DATA ANALYSIS AND HYPOTHESIS ............. 2 RESULTS: sisi58 cairn tans cere eee Oe eee 6 DISCUSSION, i577. cas Soc eee eee 6 APPENDIX 3 ..cciothe Sto ee OE 7
The use of trade, firm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the U.S. Department of Agriculture of any product or service to the exclusion of others which may be suitable.
United States Department of Agriculture
Forest Service Intermountain Forest and Range Experiment Station
Research Paper INT-281
July 1981
Monoterpenes of Lodgepole Pine Phloem as Related to Mountain Pine Beetles
Walter E. Cole, E. Park Guymon, and Chester E. Jensen
INTRODUCTION
The role of food quantity in mountain pine beetle (MPB) population dynamics in lodgepole pine is well-documented in the literature. The thickness of phloem within trees in a stand determines whether the insect can prosper there. Beetles tend to select trees that possess the thickest phloem in a stand where trees have similar diameters, and they often select that portion of an individual tree having the thickest phloem (Roe and Amman 1970). The mountain pine beetle is food-limited in those stands of lodgepole pine where developmental tempera- tures are optimum (Cole and Amman 1969). When beetles have killed most of the larger, thick-phloem trees, they are forced to attack and raise brood in the smaller residual trees. These trees have reduced capacity for supporting brood development be- cause of generally thinner phloem. Subsequently, the popula- tion declines (Cole and others 1976).
While the role of phloem quantity in beetle population dynam- ics is well documented, that of phloem quality is not. Smith (1965) has shown that vapors of the monoterpenes from west- ern white pine (Pinus ponderosa) vary in toxicity to the western pine beetle (Dendroctonus brevicomis) in the following de- scending order: limonene > A3-carene > myrcene > B-pinene ~ B-pinene > control. The monoterpene composition of oleore- sin in lodgepole pine (Pinus contorta var. muriayana) was: B-phellandrene, 69.4 percent; a-pinene, 6.4 percent; A;- carene, 8.9 percent; B-pinene, 5.7 percent; myrcene, 3.9 percent; camphene, 0.5 percent; limonene, 2.4 percent; sabinene, 2.1 percent; and a-phellandrene, 0.7 percent (Smith 1964).
In most terpene studies where a variety of pine species were considered, cortical oleoresin differed qualitatively between species but not within species. Coyne and Keith (1972) found no distinct differentiation, either qualitatively or quantitatively, between monoterpene composition of loblolly (P. taeda) and
slash (P. ellioti’) pines within or outside of known southern pine beetle outbreaks. Monoterpenes provide bases for distin- guishing host species but not for distinguishing resistant trees from check trees (Coyne and Critchfield 1974). Hanover (1975) identified an apparent genetic hierarchical regulation of the major terpene fractions in lodgepole pine. These discrete gene- tic variations may relate to pest (insect) behavioral patterns, as indicated by differing resistance levels of trees to their respec- tive pest species.
A continuing question is whether tree-to-tree differences in phloem constituents, particularly the monoterpenes, are coinci- dent with the characteristic MPB attack and survival pattern. Alpha-pinene has been the usual monoterpene used in experi- mentation with pheromones and beetle behavior. However, Moeck (1980) mentions that a-pinene is not an effective pher- omone component in lodgepole pine. While peripheral informa- tion has been developed in this study, the emphasis has been on monoterpene content of the phloem and its relation to tree characteristics previously found to be linked to MPB population dynamics.
MATERIALS AND METHODS
We took three 5.08 by 5.08 cm phloem samples at breast height from each of 86 uninfested trees distributed over 20 acres (8.1 ha) on the Cache National Forest in 1975. Trees ranged from 12.7 to 50.8 cm in diameter at breast height (d.b.h.). Samples were taken three times during the season: June 6, July 10, and July 31. The samples were transported to the laboratory and frozen on the same day they were removed from the trees. Two samples per tree were analyzed as de- scribed later, and one sample stored (frozen) as a backup sample.
The samples were stored in the lab at — 25° C. Each of the first two phloem samples were separated from the bark and ground in a Wiley grinder at 20-mesh size by freezing the sample in liquid nitrogen and by passing large amounts of dry ice through the grinder to keep the grinder cold.
Soluble nitrogen and soluble sugars were extracted with 80 percent ethanol. Insoluble products underwent chemical di- gestion in order to convert them into a soluble form that could be analyzed. Insoluble nitrogen in the sample was converted to ammonia by repeated digestion with a 20 percent sulfuric acid and cleaned with hydrogen peroxide (Hodges and others 1968).
To analyze terpenes, 0.2 to 0.3 g of the ground phloem was placed in a vial with 2 ml of isopropyl! ether (free of alcohols, chromatoquality reagent) in a sealed vial and shaken for at least 2 days. We then put 10 microliters of this solution in a Varian Aerograph series 1700 gas chromatograph with a flame ioniza- tion detector. The identification and quantity of each component was determined by running dilute standards of the pure compo- nents. The peaks were cut out, and the quantity of each compo- nent determined from its peak weight. We used a 1.83-m col- umn packed with Porapack Q because the water in the sample from the phloem did not affect this column packing. The injector temperature was 275° C, detector temperature 250° C, carrier gas (high purity helium) 40 psi, and column temperature was programmed from 50° to 250° C at 10° per minute. Ultra high purity hydrogen and air were used for hydrogen detection.
Laboratory analysis was focused on monoterpenes, soluble nitrogen, total nitrogen, reducing sugars, starches, pentoses, and hexoses. Nitrogen was analyzed by the colorimetric Nes- sler Method (Jacobs 1965). Insoluble nitrogen was determined as the difference between total nitrogen and soluble nitrogen. Sugars, hexoses, and pentoses were determined at the same time with the cysteine and sulfuric acid general reaction on carbohydrates (Dische 1955). Their absorption spectra were then read at 320 mu and 405 mu, which allows the determina- tion of both sugars. Reducing sugars were determined by methods discussed in Dische (1955). Starches were hydro- lyzed and then determined by the same procedure as the sugars.
DATA ANALYSIS
Presented in table 1 are average percentages of the phloem (by weight) and associated standard deviations found in dry matter, Sugars, starch, nitrogen, and monoterpenes for the trees sampled. Means for June 6 were compared statistically with comparable means at (a) July 10 and (b) July 31.
To establish possible links between tree characteristics and phloem constituents, the latter were fitted as linear functions of all combinations of six pertinent tree characteristics: d.b.h., percent crown length of total tree height, height, phloem thick- ness, average radial growth for the 5 years prior to sampling, and age.
Results of the regression screen are summarized in table 2 and show that rather weak regression information (R2) was developed throughout. The July 31 monoterpenes were, however, most strongly related to the tree characteristics evalu- ated. While even the strongest of these, phloem depth and growth, seem of marginal strength (0.14 < R® < 0.34), they do confirm the presence of associated linear, positive increases in monoterpenes. The results provide an information base neces-
Table 1.—Selected lodgepole pine phloem constituents, percent
by weight. i June 6 July 10 July 31 Constituent x s x s x s
soso Percent of total phloem weight ------
Dry matter O4GT N25, 45:2) 4 4a 03 -Percent of phloem dry matter weight-
Soluble pentoses 2.0 me) C0) WIG 47 102 Soluble hexoses 1e5 43° 4.3 il GS} 118} Total 3.5 ley WOKS WS KO) — 4} 74 All pentoses S37/ 98 114 272 82 1.44 All hexoses Poll dil 743 NO) NS) DG Total 64 162 19.2 3.57 19.8 3.30
Soluble reduced sugars SIS leON 222 82 1.8 82
Starch 3:0) SAROM 910) SHS 8!8) = 2:97, Insouble nitrogen ols} HD i 2 ah) 02 Total nitrogen sls) OY ike) = 08} 12 08} Monoterpenes a-pinene 052 .058 .039 .033 .030 .026 8-phellandrene 2038 277 140 124 1144 (127 3-terpenes 120 .192 .077 .067 .064 .054 (3-carene + myrcene + a-pinene) Total 375 447 .256 .203 .238 .191
sary to the development of more advanced hypotheses, to be evaluated with new data when available. In this case, an in- teractive hypothesis was developed from the July 31 data. We used two of the variables exhibiting the strongest linear effects (phloem thickness and growth) and one weak variable (d.b.h.) that proved reasonably strong in past MPB dynamics models.
Here, “total terpene” data for July 31 were partitioned over the ranges of phloem thickness, tree growth, and tree d.b.h. and were explored graphically for interactive effects. The data appeared to support a three-way interaction characterized by: positive, shallow concave-upward effects for phloem thickness and growth; a more-or-less bell-shaped effect for d.b.h., max- imizing at about 10.5 inches (26.67 cm); and convergence to zero with low growth and phloem thickness. The d.b.h. effect is not oriented at zero but is not meaningful at zero anyway. These effects were in general accord with the mountain pine beetle preference for larger, more vigorous trees, although the rather strong negative trend in terpene content for larger trees — d.b.h. > 10.5 inches (26.67 cm) — was not. Nevertheless, d.b.h. was retained in the model and the resulting four- dimensional relation was formulated mathematically using the techniques specified by Jensen (1973, 1976, 1979) and Jensen and Homeyer (1970, 1971), and was refitted to the data set from which it was partially derived, by weighted’ least squares. The final hypothesized form (R? = 0.39, Sy.x = 0.15) is shown graphically in figure 1 and mathematically in appendix table 7.
‘Variance about the initial model Y was expressed as a function of Y. The inverse of this, 1/Y*’, was used as the fitting weight.
Table 2—Summary of significant (Pr 0.05) coefficients of determination (R?) for the linear regression screens of independent variables for
three sampling dates
Date and Total independent Dry Soluble Soluble — soluble variable matter pentoses hexoses sugars JUNE 6 wen nnn nnn nnn nnn nnn nnnenne enn nnnnen nen enn nec nne nen nne enn enee Diameter at
breast height (D) Length of crown,
% of total Height Phloem thickness (P) Growth, average annual
5-yr. radial (G) 0.06 0.05 Age 0.05 .07 .06 JULY 10 Diameter at
breast height (D) Length of crown,
% of total Height Phloem thickness (P) 12 .08 Growth, average annual
5-yr. radial (G) 05 04 Age 09 05
JULY 31 Diameter at breast height (D) Length of crown, % of total .08 Height Phloem thickness (P) .06 .04 .06 Growth, average annual 5-yr. radial (G) .07 .07 Age 10
‘Total hexoses and total sugars were screened with nonsignificant results.
Terpenes
B-pinene + B-phellan- carene + Total a-pinene drene myrcene
.0 0 QQ
UU fon)
0.12 0.08 0.10 0.12 05
18 aS 09 .28 11 24 15 .07 .08 09
Additive effects
.30 lhl 25 16 .30 12 .26 18 07 .O7 .07 07
Additive effects
07 07 07 07 10 10 10 10 06 08 07 18 18 18 30 18 34 26 24 14 26 26
Additive effects
35 21 39 34 35 22 39 34
1.4 = 12 be Ga 10 a2 o = SS 08 Settle io wa zs 0.4- Sz | oo a 0.2 z 0 0 4 8 12 0 10.2 20.3 30.5 D.B.H.
5- YEAR RADIAL GROWTH IN CM
—05 12
0.3 0.76 Onli Onco
IN CM 0.20 0.51 0.15 0.38 fp 0.10 0.25 pyHioem VA THICKNESS 0.05 0.13
40. 6 50.8 CM
Figure 1. — Hypothesis: total monoterpene percentage of lodgepole pine phloem dry weight, as a function of d.b.h., phloem thickness, and average radial growth.
The July 31 monoterpene percent of phloem dry weight in trees ranged from 0.03 to 1.10 percent. Almost 40 percent of the variance (R? = 0.39) about the mean of 0.238 was explained by the regression of monoterpene percent on the strongly interact- ing independent variables, phloem thickness, growth, and tree diameter (fig. 1).
The unexpected bell-shaped effect over d.b.h. is somewhat deceptive because there is a rather strong correlation between phloem and d.b.h. The d.b.h. effect is better characterized by the monoterpene trace over the d.b.h.-phloem line of correlation (fig. 2). There it can be seen that monoterpene content reaches
amaximum at about 13 inches (33 cm) and, although the trend is slightly down thereafter, content at 20 inches (51 cm) still exceeds that for 8-inches (20 cm) trees.
Component monoterpenes were explored with much the same results as for the monoterpene sum. So, the mathemati- cal form for the sum was adopted for the components and was scaled to the data for each component using weighted (1/Y*’) least squares (fig. 3). The coefficients for component models were subsequently adjusted to equal, in sum, that for the all- component model. As a result, contents for the sum of compo- nents equal that of the all-monoterpene model at all combina- tions of d.b.h., phloem, and growth.
TOTAL TERPENE CONTENT IN PHLOEM BY WEIGHT (PERCENT)
A: Phloem = 0.0617 + 0. 00417 (d.b.h.) Sy.x 20.017, R2= 0,505
B: Terpene content trace over phloem- d. b. h. correlation
0 4 8 12 16 20 IN
30.5 40. 6 50.8 CM
D.B.H.
IN CM 0.20 0.51] Wj, 0.15 0.38 VY 0.10 0.25 PHLOEM THICKNESS 0.05 0.13 0
Figure 2. — Hypothesis: total monoterpene percentage of lodgepole pine phloem dry weight, trace over d.b.h./phloem correlation at average annual radial growth (0.159 inches, 0.404 cm).
2 hg
oO
as
ae
Qe 43
Bo Kian
Sx
So
SF 02
w Gs
ae
ea (ee
oe Ol
= 6
ey
&
- 9
0 r 8 12 16 0 10.2 20.3 30.5 40. 6 D.B.H.
20 ~=IN 50.8 CM
Figure 3. — Hypothesis: total monoterpene percentage, of lodgepole pine phloem dry weight over d.b.h. and phloem thickness at average annual radial growth (0.159 inches, 0.404 cm).
RESULTS
Means and associated standard deviations are shown in table 1 for all phloem contents evaluated in this study. Numbers of sample trees varied from 79 to 86 depending on date of sample and phloem component analyzed.
Means for July 10 and 31 were consistently lower than those of June 6 for dry matter, soluble reducing sugars, nitrogen, and monoterpenes. Differences were significant (Pr < 0.05) for the first three components and less so for the monoterpenes (0.10 < Pr < 0.05). Starches and other sugars showed increases from June 6 (Pr < 0.05).
We found monoterpene percentage of the phloem dry weight to be extremely small (0.238) and, of this, individual monoterpenes were distributed in about the same proportion as found by Smith (1964) in “pure” oleoresin (table 3). And, in either case, 8-phellandrene is by far the largest monoterpene component, followed by the 3-terpene group and a-pinene, respectively. Note that the averages are greater in larger trees with thicker phloem (and vice versa) according to the interactive hypothesis (fig. 1). Too, the expected trend over the d.b.h.- phloem trace (fig. 2) increases to a peak at about 13 inches (33 cm) d.b.h., decreasing thereafter to a low at 20 inches (51 cm) comparable to that at about 9 inches (23 cm).
Note that the percentages of phloem dry weight reported in table 1 and in figures 1 through 3 are based on the monoterpenes measured in this study only. A small reduction in percentages for components could be expected with upward adjustment of the monoterpene sum by 6 percent, to achieve comparability to Smith’s (1964) percentages (table 3).
Table 3.—Proportional distribution of monoterpenes: ‘“‘pure”’ oleoresin versus phloem dry matter
In phloem In ‘“‘pure” dry matter oleoresin this study, Monoterpenes (Smith 1964) 7/31/75" noncnonernnnnnn== Percent by weight ---------------- a-pinene 6.4 11.9 8-phellandrene 69.4 57.0 3-terpenes 18.5 25.4 (3-carene + myrcene + a-pinene) Others 5.7, (5.7) (camphene + limonene + sabinene + a-phellandrene) Total 100.0 100.0
‘Original percentage adjusted for 5.7 percent of “others” not evaluated.
DISCUSSION
The inference limitations in this study are rather servere because the sample trees involved are from a single, infinitely small stand relative to the whole. But in the absence of stronger information on lodgepole pine phloem, our findings provide a data-base opportunity to develop hypotheses for more exten- sive study.
Table 1 contains mean percentages and standard deviations for a variety of phloem components, all of which are likely to have some impact on MPB population dynamics. The data on sugars, starch, and nitrogen are simply documented here for general interest. We note, however, that most sugars and starch are at low levels in the spring and that soluble reducing sugars, nitrogen, and monoterpenes are relatively high. These trends follow expectations based on seasonal tree physiology, but because all but monoterpene relations to tree characteris- tics appeared to be extremely weak (table 2), we did not attempt to develop such information further.
Respective (but very low) concentrations of monoterpenes in the phloem are parallel in proportions of the monoterpene sum to those found by Smith (1964) in pure oleoresin produced in lodgepole pine (table 3). Beta-phellandrene in both studies proved to be, by far, the largest component of the monoter- penes, and so might easily have the greatest impact on MPB activities. We note that while a-pinene has been found to be an effective pheromone in western white pine, itis not for lodgepole (Moeck 1980).
But whether it is B-phellandrene or some lesser component of the monoterpenes, concentrations in the phloem appear from the hypothesis developed (fig. 2), to increase with tree vigor and size, up to an optimum d.b.h. of about 13 inches (33 cm).
It has been established from past research that threshold diameters in lodgepole pine for successful MPB reproduction are generally in the 8-inch (20-cm) to 9-inch (23-cm) range. And reproduction success is known to be high in larger, more vigo- rous trees. This information, together with the coincidence of relatively high monoterpene content for larger trees (9-inches [23-cm] to 20-inches [51cm] d.b.h.; see fig. 3), is perhaps sug- gestive of an attractant role for any one or all of the monoter- penes. It would also appear that monoterpene toxicity levels studies by Smith (1965) are apparently not being reached in the phloem, based on the level of MPB success in larger trees.
The hypothesis developed in this study (fig. 1-3 and appen- dix) should help to identify points of future study emphasis and may be rescaled (as a unit) and evaluated for performance on new data sets (Jensen 1979).
APPENDIX
Table 4.—Hypothesis values for figure 1. Monoterpene percentage of lodgepole pine phloem dry weight. All-monoterpene % = 1.10301 * (model)'
D.b.h. Average annual Phloem 4 8 10.5 12 16 20 (inches) radial growth thickness 10.2 20.3 26.7 30.5 40.6 50.8 (cm)
Inches cm Inches cm 0.1 0.25 0.05 0.13
10 25
15 .38 j : : j :
.20 51 PSG 2or S80 V769N-597- 418 3 16 .05 13 : A A722
10 25 : A .356 ;
aS .38 .330 .627 .700 .668 .518 .363
.20 ail 516 .979 1.094 1.043 .810 .567 AS WRI, 05 as} OSI b4S 72s 1641127, 089
10 25 .238 .504 .480 .373 .261
15 38 446 846 699 .490
.20 51 .695 1.321 1.476 1.407 1.092 .765
'The enclosed areas are represented by one or more data points. The same is true for the monoterpene component tables that follow.
Table 5.—Hypothesis values for monoterpene components, percentage of lodgepole pine phloem dry weight (no related figure in text)
D.b.h. Average annual Phloem 4 8 10:5" 2 16 20 (inches) radial growth thickness 10.2 20.3 26.7 30.5 40.6 508 (cm) Inches cm Inches cm a-pinene % = 0.15787 * (model) 0.1 0.25 0.05 0.13 0.007 0.012 0.014 0.013 0.010 0.007 10 25 .019 .036 .040 .038 .029 .020 15 38 .034 .066 .074 .070 .055 .038 .20 coil 0055 .014 115 .110 .085 .060 3 76 .05 sls} .008 .016 .018 .017 .014 .009 10 25 025 .048 .053 .051 .040 .028 iS 38 .048 .090 .100 .096 .074 .052 .20 51 1074 139) 157 149) 116 <081 5 1.27 05 13 {Oil O22) 2025 023 ee 0118s 2 Os 10 25 034 .064 .072 .069 .053 .037 3S 38 (0645 1122) 5135) 2129-1100). 2070 .20 51 100 .189 .211 .201 .156 .109
B-phellandrene % = 0.64292 * (model)
0.1 0.25 0.05 0.13 0.026 0.049 0.055 0.052 0.041 0.028 10 .25 075 144 N61 53 1119) 2083
Bilis) 38 M42" 7270 2301 4287-223) 356
.20 51 .221 420 .470 .449 .348 .244
3 76 05 3 035 .066 .074 .071 .055 .039 10 29 102 .194 .218 .208 161 .113
AS 38 a192> 2365) -408" =389) 23025 2252
.20 Zoi 300 .571 .637 .608 .472 .330
25 1.27 05 13 047 .090 .100 .096 .074 .052 .10 229) 4138) 2263) 22938)" 280) i217 ali52
“15 38 .260 .493 .551 .525 .408 .285
.20 x) 406 .770 .860 .820 .637 .446
(B-pinene + 3-carene + myrcene) % = 0.30222 * (model)
0.1 0.25 0.05 0.13 0.012 0.023 0.026 0.025 0.019 0.013 10 25 035 .068 .075 .072 .056 .039
ails) 38 067 .126 .141 .135 .105 .073
.20 foil N05) 4198) 2221 22h 64s wea
3 76 .05 13 017 .032 .035 .033 .026 .018 10 25 .049 .091 .102 .098 .076 .053
alls) 38 090 .172 .192 .183 .142 .099
.20 SoH 141 .269 300 .286 .222 .155
2) 127, 05 13 022 .043 .047 .045 0385 .024 10 25 10665 4124" 139)" -132) 02) 072
SUS) 38 122 .231 .259 .247 192 .134
.20 51 191 .362 .405 .385 .299 . .210
Component
Table 6.—Hypothesis values for figures 2 and 3. Monoterpene percentage of lodge-
pole pine dry weight, at average annual radial growth = 0.159 inches (0.404 cm), average for 86 trees. Percent by weight of phloem at average annual
radial growth = 0.159 inches (0.404 cm)
20 (inches)
Table 7.—Mathematical descriptors for figures 1-3 and appendix tables 4, 5, and 6.
Monoterpene Models
Percent monoterpene content = (21.0621 * YPP * P'®°®) * K; (Dice 18'S): jaca 2
For D < 10.5 19 0.395 YPP = YPD * (1.00165 *e — 0.00165) (GSiS he D) eee For D > 10.5 A 28 0.49 YPP = YPD* (1.07092 *e — 0.07092) For0 <D S 22 YPD = 0.38 + 1.0292 * G'* S R? ess K, = 1.10301, all monoterpenes 0.387 0.150 Kz = 0.64292, B-phellandrene 0.402 0.098 K3 = 0.30222, B-phinene + 3-carene + myrcene 0.352 0.044 K, = 0.15787, a-pinene 0.213 0.023 where
P = phloem thickness, inches; D = tree d.b.h., inches; = average annual radial growth, last 5 years, inches.
(@) |
‘Conservative estimates.
(cm)
Ne A am DED Ne” eh va ate) Phloem 4 8 105 12 16 thickness 10.2 20.3 26.7 30.5 40.6 50.8 Inches cm a-pinene 0.05 0.13 0.007 0.013 0.015 0.014 0.011 0.008 10 25 020 .038 .043 .041 .032 .022 15 .38 .038 .072 .081 .077 .060 .042 .20 2511 1059) iS A267 1120"): 093)" 065 8-phellandrene 05 a3 .028 .053 .060 .057 .044 .031 10 .25 10825 FN STAF NT 5N GHP E29 091 aS) 38 b> Ee G293E RS2Bmer Sili2im=24ayn AAO .20 51 242 .458 .512 .488 .379 .265 B-pinene + .05 ac} 013° 1025. 028) 027 <_.02)\ .015 3-carene + 10 25 039 .074 .082 .078 .061 .043 myrcene alts) 38 073 1388 .154 .147 .114 .080 .20 {5H Aen) 24, e2eON aliS, yale All terpenes .05 13 .048 .092 .102 .098 .076 .053 (sum of those 10 25 142 .269 .300 .286 .222 .156 IS) 38 2265) “£5044 W563) 7:536) 1-416) 2292 .20 51 .414 .786 .879 .837 .650 .455
PUBLICATIONS CITED
Cole, W.E., and G. D. Amman.
1969. Mountain pine beetle infestations in relation to lodge- pole pine diameters. USDA For. Serv. Res. Note INT-95, 7 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Cole, W. E., G.D. Amman, and C. E. Jensen.
1976. Mathematical models for the mountain pine beetle-- lodgepole pine interaction. Environment. Entomol. 5(1):11- 19.
Coyne, J. F., and W. B. Critchfield.
1974. Identity and terpene composition of Honduran pines attacked by the bark beetle Dendroctonus frontalis (Scoly- tidae). Turrialba 24(3):327-331.
Coyne, J. F., and G. C. Keith.
1972. Geographical survey of monoterpenes in loblolly and shortleaf pines. USDA For. Serv. Res. Pap. SO-79, 12 p. South. For. Exp. Stn., New Orleans, La.
Dische, Z.
1955. New color reactions for determination of sugars in polysaccharides. /n Methods of biological analysis, vol. Il. p. 323-327. Interscience Publishers, Inc., New York.
Hanover, J. W.
1975. Comparative physiology of eastern and western white pines: oleoresin composition and viscosity. For. Sci. 21(3):214-221.
Hodges, J. D., S. J. Barras, and J. K. Mauldin.
1968. Amino acids in inner bark of loblolly pine as affected by the southern pine beetle and associated microorganisms. Can. J. Bot. 46:1467-1472.
Jacobs, S.
1965. The determination of nitrogen in biological material. /n Methods of biochemical analysis, vol. XIII. p. 251-252. Interscience Publishers, Inc., New York.
Jensen, C. E.
1973. Matchacurve-3, multiple-component and multi- dimensional mathematical models for natural resource models. USDA For. Serv. Res. Pap. INT-146, 42 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Jensen, C. E.
1976. Matchacurve-4, segmented mathematical descriptors for asymetric curve forms. USDA For. Serv. Res. Pap. INT-182, 16 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Jensen, C. E.
1979. e*, a function for the modeler. USDA For. Serv. Res. Pap. INT-240, 9 p. Intermt. For. and Range Exp. Sin., Ogden, Utah.
Jensen, C. E., and J. W. Homeyer.
1970. Matchacurve-1 for algebraic transforms to describe sigmoid- or bell-shaped curves. 22 p. USDA For. Serv., Intermt. For. and Range Exp. Stn., Ogden, Utah.
Jensen, C. E., and J. W. Homeyer.
1971. Matchacurve-2 for algebraic transforms to describe curves of the class X". USDA For. Serv. Res. Pap. INT-106, 39 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Moeck, H. A.
1980. Field test of Swedish ‘drainpipe’ pheromone trap with
mountain pine beetle. Bi-mon. Res. Notes 36(1):2-3. Roe, A. L., and G. D. Amman.
1970. The mountain pine beetle in lodgepole pine forests. USDA For. Serv. Res. Pap. INT-71, 23 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Smith, R. H.
1964. The monoterpenes of lodgepole pine oleoresin. Phy-
tochemistry 3:259-262. Smith, R. H.
1965. Effect of monoterpene vapors on the western pine
beetle. J. Econ. Entomol. 58(3):509-510. Smith, R. H.
1975. Formula for describing effect of insect and host tree factors on resistance to western pine beetle attack. J. Econ. Entomol. 68(6):841-844.
Cole, Walter E., E. Park Guymon, and Chester E. Jensen.
1981. Monoterpenes of lodgepole pine phloem as related to mountain pine beetles. USDA For. Serv. Res. Pap. INT-281, 10 p. Intermt. For. and Range Exp. Stn., Ogden, Utah 84401.
Phloem samples taken from 86 healthy lodgepole pine trees were analyzed for content of dry matter, starch, various forms of sugar and nitrogen, and of selected monoterpenes. B-phellandrene was, by far, the most prevalent of the mono- terpenes. An interactive hypothesis is developed for terpene content as a function of phloem thickness, radial growth, and tree diameter. Here, high concentrations of monoterpenes coincide with upper tree diameters, the expected region of high mountain pine beetle survival success.
KEYWORDS: phloem constituents, lodgepole pine, Pinus contorta var. latifolia, sugars, starch, nitrogen, monoterpenes, mountain pine beetle, De- ndroctonus ponderosae.
WY U.S. GOVERNMENT PRINTING OFFICE:
1981-0-780-811
The Intermountain Station, headquartered in Ogden, Utah, is one of eight regional experiment stations charged with providing scientific knowledge to help resource managers meet human needs and protect forest and range ecosystems.
The Intermountain Station includes the States of Montana, Idaho, Utah, Nevada, and western Wyoming. About 273 million acres, or 85 percent, of the land area in the Station territory are classified as forest and rangeland. These lands include grasslands, deserts, shrublands, alpine areas, and well-stocked forests. They supply fiber for forest in- dustries; minerals for energy and industrial development; and water for domestic and industrial consumption. They also provide recreation opportunities for millions of visitors each year.
Field programs and research work units of the Station are maintained in:
Boise, Idaho
Bozeman, Montana (in cooperation with Montana State University)
Logan, Utah (in cooperation with Utah State University)
Missoula, Montana (in cooperation with the University of Montana)
Moscow, Idaho (in cooperation with the Univer- sity of Idaho)
Provo, Utah (in cooperation with Brigham Young University)
Reno, Nevada (in cooperation with the University of Nevada)