· Reprinted from the JOURNAL OF FORESTRY, Vol. 62, No. 4,' April 1964 Purchased by the U.S. Forest Service for officiall.lse. Simultaneous Variations in Seasonal Height and Radial Growth-of Young Douglas-Fir Edward J. Dimock II RELATIONSHIPS between seasonal in­ crements in height and diameter of forest trees have long stirred more than casual interest among plant scientists (7, gy. It has been found that bud bursting generally shows a . close correlation with' cambial re­ activation following' winter dor­ mancy. However, the physiologi­ eal relationship between apical and cambial growth J.:emains obscure. and leaves much for further study. To date; the bulk of published find­ ings stems from study of seedlings , and saplings. This paper reports within-season changes in diameter and height on Douglas-fir trees ex­ ceeding 30 feet tall. These changes, recorded s i m ult a n e o u s l y, were charted throughout two growing seasons to elucidate grosser fea­ tures of the height-diameter rela­ . tionshi:r>' Tree Description Ten. dominimt- Douglas-fir trees. were selected for measurement. At ' time of first observation during the spring of 1957, these trees averaged 16 years in total age, 6.03 THE AUTHOR is research forester on the .staff of the Pacific Northwest Forest and Range Expt. Sta., Forest Service, tr. S. Dept. Agric., Portland, Ore. Abstract. Progressive changes in bole circumference and total height were simultaneously measured in 10 Douglas-fir trees during the 1957 and 1958 grow­ ing seasons.· At the start of each season, substantial increases in circumference 'began by mid-April and preceded comparable increases in height by approxi­ mately one month. Swelling or bursting of terminal buds did not seem to be 1m essential precursor to radial growth. Height growth was mostly complete by August, whereas' apparent radial increments were still evident during October and November. illches in breast-high diameter, and 32.6 feet in total height. They formed part of an open stand that originated naturally on an old field site of medium quality near McCleary, Wash. All trees were growing rapidly and possessed the smooth bark and long' terminal shoots characteristic of their age and thrift. They were sufficiently separated to minimize direct com­ petition for growing space. Meas­ urements during the growing sea­ sons of 1957 and 1958 showed re­ spective annual increases averag­ ing 1.006 and 1.336 i n c h e s in breast-high circumference, and 2.69 and 2.73 feet in total height (Ta­ ble 1). Tree nleasurement Simultaneous determinations of circumference and height were re­ corded weekly early in each seaso'n; 4. TABLE I.-ANNUAL INOREASES IN HEIGHT AND CIROUMFERENOE OF DOUGLAS-FIR TREES DURING THE 1957 AND 1958 GROWING SEASONS Tree no. I 2 3 4a 4b1 5 6 F 7 8 r 9 10 \ Initial measurements, spring of 1957 Age D.h.h. Height Years Inches Feet 17 17 15 15 17 16 22 13 13 14 14 6.22 5.12 5.46 6.49 6.84 6.97 7.56 5.60 5.75 5.90 5.27 34.8 32.2 30.3 ' 31.7 34.5 31.1 46.0 28.9 31.3 32.3 27.8 Annual circumferential increase 1957 1958 ' -Inches -- . - 0.860 1.180 .465 1.000 .960 1.140 .960 .805 1.275 1.425 Annual terminal increase 1957 -- 1958 Feet-- 1.230 1.550 1.015 3.24 2.91 1.24 2.51 3.04 2.50 .945 1.600 1.025 1.285 1.350 1.740 1.610 3.01 2.20 2.69 2.83 3.17 2.93 3.06 2.70 2..02 2.85 2.72 3.42 2.53 1Tree substituted in 1958 for tree' No. 4a whieh sustained a broken terminal shoot during the· 1957 grow.ing season. Age, d.b.h., and height are for spring of 1958. APRIL 1964 then, biweekly as seasonal incre­ ment rates began to drop. Meas­ urements were not made over the late fall and winter months, al­ though small amounts of radial stem change (7.2 percent in 1957 and 2.8 percent in 1958) occurred. All measurable height growth, how­ ever, fell within each spring-to­ fall period. Ohanges in c i r cmuf e r e n c e at breast height were determined by vernier tree-g'rowth bands of the type described by Hall (6) and recently modified by Liming (12) and Mesavage and Smith (.13). These dendrometers, custom fitted to each tree, were constructed of %-inch-wide aluminum tape such as used for numbered tree tags. Measurement and vernier scales were scribed by hand and cali­ brated to indicate circumferential changes to the nearest 0.005 inch. Oadmium-plated s t e e I springs, which held the bands in place, proved satisfactory throughout two years of use. Since tree t o p s w eI' e beyond reach, a surveyor's transit was used to determine height changes. Two permanent hubs, each marked by a finishing nail driven into the top of a firmly anchored stake, served for all 10 trees; 4 trees were measured from one hub, 6 from the other. Thus, although instrument height varied slightly, orientation and horizontal distance from hub to tree remained identical at each measurement. Horizontal distances between each hub and the bole axes of trees common to it ranged from 44.5 to 82.0 feet. The observer read two vertical angles to the nea.rest minute for each tree-'--all as deflections from horizontal: (1) a base angle for fixed reference to the top edge of each tree's growth band and (2) an angle to the tree top. Periodic height changes were determined by comparing total heights calcu­ lated separately for each observa tion date in the following manner: height = (h - til)d + 4.5 where h = tangent of angle to tree top where t2 = tangent of angle to top edge of growth band where d = horizontal distance to tree in feet. 253' " o , 1958 1.25 w u :z: w '" 1.00 1957 §j o 0.75 W to :z: "'" :I: U W > 0,50 § :0 § 0.25 U ..J "'" JUNE ---- oYr ---- -- AUG. -- 175 145 205 SEPT. OCT. L----,-L---. 235 NOV. ----,- 265 __ 325 NUMBER of DAYS since b gj ning of each year FIG. I.-Relationship of change in circumference (b.h.) to time for 10 Douglas-fir trees during 1957 and 1958 growing seasons. 1'----'---, 3.00,, 1958 1957 o =- 2.50 to iii ::z: 2.00 0; W to :z: "'" {51.50 W > ;;; :3 ],00 '"' :0 U ..J 0.50 :z: :z: "'" APRIL 85 AUG. , 115 145 175 NUMBER of DAYS since beginning of each year 205 1 I SEPT. 235 OCT. I 265 NOV. 295 I 325 . FIG. 2.-Relationship of change in total height to time for 10 Douglas-fir trees during 1957 and 1958. growing seasons. loo �f------t_-��::�����---� 80 ..J 60 W ;" :z: o i)l 40 o 20o ------iL---�--+--� o oL-, APRIL 85 l1S JUNE MAY I 145 ---- HUMBER of DAYS since beginning of each yeor 175 ULY I --+_ J I 205 AUG. I - SEPT. -L- 265 235 ' OCT. - NOV. r._ -- 325 295 . ' FIG. 8.--Oumulative change in .circumference (b.h.) and in total height for 10 Douglas-fir trees in relation to time (av rage for combilied 1957 and 1958 growing ' seasons). 254 JOURNAL OF FORESTRY Seasonal Growth MeasuraJ:>le circumferential in­ creases began at different times during the 1957 and 1958 growing seasons (Fig. 1). In 1957, detect­ able increases began in mid-April. However, in 1958, all trees were increasing during the first measure­ ment interval, and it appears likely t11at radial growth began shortly before the first 0 b s e r v a t i o n on March 28. Growth rates during both years, though lower in 1957 than in 1958, remained fairly constant through mid-July. Then, while 1957 growth generally leveled off gradually" in 1958 there was an abrupt and prolonged period of circumferential shrinkage, and ex­ pansion did not resume until late August. In both years, a' slow increase continued through the last observation period, which ended on October 25, 1957 and on November 21, 1958. Seasonal p a t t el' n s of height growth showed much greater simil. arity between the two observed years. Detectable increases, due to swelling of a few terminal buds, began in both years during the latter half of April (Fig. 2). How­ ever, most terminal buds did not burst until nearly mid-May, and significant increments in height did not begin until then. The terminals of all trees lengthened rapidly until early August, when growth quickly leveled off in a manner fairly' consistent for both seasons. Height growth was 95 percent com­ plete by early August 1957 and by late JUly 1958. Erratic meas­ urements were responsible for ap­ parent slight decreases in height, which appeareq. once in 1957 (Sep­ tember) and twice in 1958 (April and August). Major growth trends in both radius and height were rehitivel;Y consistent for all trees within the 10-tree sample. For example, the shrinkage in circumference during the 6-week interval from July 18 to August 29, 1958 was based on changes Over three 2-week inter­ vals. Among 30 individual circum­ ference measurements, 26 showed decrease, 2 no change, and 2 a slight increase. At both ends of the major growth periods, how­ ever, deviations of individual trees significantly affected averages of height and circumference. To ilhis­ trate, one tree, by rapidly straight­ ening a'leaning terminal shoot be­ tween September 27 and October 11, 1957, increased 0.48 foot in height-eight times as much change as shown by any other tree during the same period. Cumulative changes in height and circumference described a sig­ moid-curved relationship with time (Figs. 1 and- 2). Height curves tended to sho'w l' a t h e l' abrupt changes in rate both preceding and following the major growth period. Changes in circumference showed a similarly abrupt increase leading into the period of rapid growt.h, but seasonal reduction in rate was' gradual and somewhat erratie These general trends are more clearly illustrated by combining data for the 1957 and 1958 sea­ sons and e x p r e s s i n g periodic changes as proportional to total change (Fig. 3). Though such a procedure masks significant be­ tween-season differences, stemming from varying climatic patterns, it defines trends that are probably closer to long-term averages. From the combined data it is evident that substantial increases in height did not begin' until about May 10fully one month after the start of comparable increases in circumfer­ ence. Height growth then pro­ ceeded at a faster rate than eircum­ ference growth until both reached 80-percent completion by JUly 10. Height growth reached its maxi­ mum for the season shortly there­ after, while the curve of circum­ ferential growth was still ascend­ ing gradually and nearly 6 percent short of its maximum. Measurement of height changes by transit proved to be a workable means for determining gross trends. However, the transit did not have precision commensurate with the dendrometer. During the ea:r;ly weeks of rap. id shoot elongation, a number of terminals leaned or bent sufficient­ ly to cause serious underestimation of actual height growth. This effect was reversed later as the shoots regained their turgidity and grad­ ually straightened. Much of the apparent height growth fter early August stemmed from this mechan.; ical straightening, and not from actual. cell divisions or elongation. Discussion The apparent initiation of height growth after radial growth is a noteworthy departure from trends suggested by other workers (8,10, 15,16,.17). In most instances, bud bursting and termhial 'elongation have been observed to precede the start of radial growth 01" to occur nearly simultaneously with it. Tho downward translocation of hor­ monal substances produced by ac­ tivity of the apical iner'istems is reputed to trigger the cambial' ae­ tivity necessary for radial growth (1, 4, 5, 11, 14, 18). This theory cau,uot be refuted on the' basis of gross ob,servations reported in this study. Hpweve,r, the possibi, lity is raised that hormones necessary to stimulate radial growth of Doug­ las-fir Juay be translocated through­ out the cambium conside ably be­ fore the initiation of any obvious growth in vegetative terminals. Moreover, as is typical with the spe­ cies, reproductive buds' oc urring on several of the study trees burst shortly before mid-April in both 1957 and 1958. The appearance of strobiles fully one month before bursting of vegetative buds was more nearly coincident with the ap­ parent initiation of radial growth. In any case, many physiological processes undoubtedly operate in initiating cambial activity, and in Douglas-fir the bursting of vegeta. , r to be tive buds does not appea an essential stimulus. Though dendrometers ,accurate­ ly measure external fluctuations (2), each radial change combines the :llet effect of both swelling due to cambial growth and of shrink­ ' age or sweliing due to moisture changes within the t ee (3). Hy­ dration levels in both living and dead tissues may vary sufficiently to caUSe considerable over- or un­ derestimation of growth by en­ largement of cambial initials or by cell division. Though this effect is minimized during periods of most rapid growth, it can mask actual growth trends when cambial activity is slight. The radial growth initiation dates observed in this study may .APRIL 1964 precede the start of actual growth because radial swellings induced by accelerated moisture uptake from the soil would appear similar to those resulting from early sea­ son cambial activity. In view of climatic conditions in the Pacific Northwest, however, it is felt that hydration levels remain sufficiently high during the dormant months .' to minimize swelling due to' re­ activating sap flow in late winter, and that earliest recorded increases reasonably approximate the start of actual growth. Circumferential shrinkages oc­ curring later· in the growing sea­ son, particularly at the time of normally decelerated cambial ac­ tivity, suggest thE) more prollounced effect of dehydration as it affects apparent growth. The 6.week pe­ riod of "negative" circumferential growth in 1958, occasioned by a sustained period of rainless weath­ er, shows that dehydration can lfiore than offset. any existing low­ level cambial activity. Further. more, rehydration of living and dead tissues with the onset of late ' ummer or fall rains undoubtedly exaggerates whatever real tissue 255 building is taking place in the cam­ bium at that time. Hence, the actual cessation of seasonal radial growth can only be inferred from data in this study, and probably could no have been reliably de­ tected short of histological exam­ ination.. Literature Cited 8. 9. 10. 11. 1. BORMANN, ]'. H. 1962. Root graft­ 2. 3. 4. 5. 6. 7. ing and non-competitive relationships between trees. In: Tree growth. Ed. by '1'heodore T. Kozlowski. 'fhe Ronald Press Co., New York. Pp. 237-246. . ' ---, and T. T. KOZLOWSKI. ] 902. Measurements of tree growth with dial gage. dendrometers and vernier tree ring bands. Ecology 43 : 289-294. FRASER, DONALD A. 1956. Ecologi· cal studies of forest trees at Chalk River, Ontario, Canada. II. Ecologi· cal conditions and radial increment. Ecology 37: 777-789. . 1958. Growth meclmn· isms in hardwoods. Pulp & Paper Mag. Canada 59(10) :202-209. GOUWENTAK, C. A. 1941. Cambial activity as dependent on the pres­ ence of growth hormone and the non-resting conditionof stems. Amsterdam Acad. Sci. Proc. 44:654663. HALL, RALPH· C. 1944. A vernier tree-growth band. Jour. l!'orestry 42:742-743. KOZLOWSKI, THEODORE T. [Ed. ] . 1962. Tree growth. Tlie Ronald Press Co., New York. 442 pp. 12. 13. 14. 15. 16. 17. 18. 1962. Photosynthesis, climate, and tree growth. In: Tree growth. Ed. by Theodore T. Koz­ lowsld. The Ronald Press Co., New York. Pp. 149-164. KRAMER, PAUL J., and THEODORE T. 1960. Physiology of KOZLOWSKI. trees. McGraw-Hill Book Co., New York. 642 PT>. ·LAl>EFOGED, KJELD. 1952. The pe­ riodicity of wood formation. Danske Vidensk. Selsk. BioI. Skr. 7(3) :1-98. LARSON, PHILIP R. 1962. Auxin gradients and the regulation of cam­ bial activity. In: Tree growth. Ed: '1'he by Theodore '1'. Kozlowski. Ronald Press Co., New York. Pp.­ 97·117. LIMING, FRANKLIN G. 1957. Home-' made dendrometers. Jour. Forestry 55:575-577. MESAVAGE, CLEMEN'l', and W. S. SMITH. 1960. 'rimesavers for instaU­ ing dendrometer bands. Jour. For. estry 58:396. SAMISH, R. M. 1954. Dormancy in Anll. Rev. PlaI).t woody plants. Physiol. 5: 183-204. WAREING, P. F. 1951. Growth s tUdies. in woody species. IV. '1'he initiation of cambial activity in ring-porous species. Physiol. Plant. 4: 546-562. . 1958. The physiology J our. lust. of cambial activity. Wood Sci. 1: 34-42. WIWOX, HUGH. 1962. Cambial growth characteristics. In: Tree growth. Ed. by Theodore '1'. Koz­ lowski. '1'he Ronald Press Co., New York. Pp. 57-88. WOR , D. J. 1962. Physiology of cambial activity. In: 'rree growth. Ed. by Theodore T. Kozlowski. The Ronald Press Co., .New York. Pp. 89-95. 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