419 Effects of annual crown pruning and serial propagation on rooting of stem cuttings from Douglas-fir DoNALD L. CorEs Pacific Northwest Forest and Range Experimem Station, United States Departmem of Agriculture, Forest Service, Corvallis, OR, U.S.A. 9733/ Received Octo per 28, J 982' Accepted January 19, 1983 COPES, D. L. 1983. Effects of annual crown pruning and serial propagation on rooting of stem cuttings from Douglas-fir. Can. J. For. Res. 13: 419-424. Success of rooting from stem cuttings of 14 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) clones increased signifi­ cantly from 42% in 1974 to 67% in 1981. The increase was slightly more than 3% each year despite ortet age increasing from I 0 and 13 years in 1974 to 17 and 20 years in 1981. The greatest increase occurred from the 3rd through the 6th year of annual rooting trials. Yearly pruning of ortets and rooted ramets of the same clones is thought to have reinvigorated shoots from the pruned crowns of trees. Several annual prunings seemed to be required before average rooting potential increased markedly. No translocation of the reinvigoration stimulus from the pruned to the adjacent unpruned areas of the same trees was detected. Rooting of cuttings from ortets from pruned areas was at least 9% more than cuttings from unpruned areas. No additional invigoration from serial propagation of cuttings was noted for rooting potential, but such cuttings grew a little taller and slightly less plagiotropic 1 year after rooting than were ramets propagated from pruned ortets. Significant clonal variation was found with all treatments. Evidence of a change in physiological vigor of the ortcts and rooted ramets was externally visible in the formation of abundant lammas shoots on pruned areas of the 17- and 20-year-old ortets and rooted ramets. Abundant lammas growth is normally found only on Douglas-fir trees less than 10 years old. COPES, D. L. 1983. Effects of annual crown pruning and serial propagation on rooting of stem cuttings from Douglas-fir. C an J. For. Res. 13:419-424. . La reussite d'enracinement des boutures de rameaux de 14 clones de Pseudotsuga menziesii (Mirb. ) Franco augmenta de fa!fon nette, pussant de 42% en 1974 a 67% en 1981. Cctte augmentation se chiffrait a un peu plus de 3% par annee malgre !'augmentation de !'age des pieds-meres qui passait de 10 a 13 ans en 1974, puis de 17 a 20 ans en 1981. La plus grande augmentation dans l'enracinemcnt se produisit a partir de Ia 3c jusqu'a Ia 6" annee des essais annuels. Une taillc annuelle des pied-meres et des rameaux enracincs d'un memc clone semble avoir revigorc les ramcaux preleves des cimes emondees de ces arbres. II semble egalcmcnt que plusieurs taillcs annuclles soicnt requiscs pour obtenir un potentiel d'enracinement nettement augmente. De plus, !'auteur ne decela pas, sur un meme arbre, de transfer! de ce stimulus dans l'enracinement des parties de cime emondees vcrs cellcs voisines qui ne lc furent pas. La rhizogencse des boutures en provenance des pieds-meres aux parties de cime tai!lees s'est avcrec 9% superieure a celle des boutures prclevees sur des cimes demeurees sans traitements. De plus, !'auteur n'obscrva pas de vigueur additionnclle dans l'enracinement, suite a une propagation en serie des boutures, cepcndant de telles bouturcs pousserent un peu plus ct furent scnsiblement mains plagiotropes, I an apres l'enracinement, compara­ tivcment aux rameaux prelcves des pieds-meres emondes. On note une nette variation clonalc entre tous les traitements, de meme que !'evidence d'un changement en vigueur physiologique chez ces pieds-meres et les boutures racinees, cette derniere se traduisant par Ia formation d'abondantes pousses d'aoOt, produites sur ces parties taillees des pieds-mercs ages de 17 et 20 ans ct sur lcs bouturcs racinces. La presence d'abondantes pousscs d'aoOt represente un phenomene normal sur les Pseudotsuga ages de moins de 10 ans. [Traduit par le journal] Introduction Many reports on various rooting studies with Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) are available. The sample of existing literature presented below merely indicates the scope of past work. Propagation by stem cutting was started in the United States in the 1930's (Griffith 1940; McCoulloch 1943), but the rooting techniques were little used until the intensive interest in tree improvement in the 1960's resulted in numerous studies on rooting of Douglas-fir 'Revised manuscript received January 18, 1983. stem cuttings. Investigations were made in the fol­ lowing areas: cultural treatment (Black 1972; Cornu 1973; Ross 1975), paired cutting technique (Brix and Barker 1973), time of year (Roberts and Fuchigami 1973), bud and cambial activity (Sheila and Roberts i 975), aging (Roberts and Moeller 1978), shoot origin (Black 1972), photoperiod ( Bhella and Roberts 1974), root anatomy (Heaman and Owens 1972), rooting me­ dia (Copes 1977), growth rate and tree form (Copes 1976), and bud dormancy (Roberts eta/. 1974). Root­ ing trials in several of these areas were also made by Brix and Barker ( 1973), with shoots from adult trees. Literature on conifers generally indicates that most CAN. 420 J. FOR. RES. VOL. 1.1. llJH.1 conifers root easily when young. but rooting potential decreases as trees mature (Hyun 1967). No decrease was noted for Douglas-fir cuttings between ages I and 9 years, but a sharp decline occurred after age 9 (Black 1973). In another study, no decrease in rooting poten­ tial was noted when 14 Douglas-fir clones were tested each year from year 12 through year 16 (Roberts and Moeller 1978). The neg<ltive effect of advanced phys­ iological age of the ortet was demonstrated in France, where 64o/r of the cuttings from I 0-year-old Douglas-fir trees rooted, but only 18% of the cuttings from 60-year-olcl trees rooted (Cornu 1973). A del'inite de­ crease in rooting potential occurs as Douglas-fir trees mature, but the literature is not clear about when the decrease occurs. It does not appear to be correlated with the onset of flowering (Roberts and Moeller 1978). Annual pruning or hedging treatments ha vc been suc­ cessfully used in "Radiata pine" to retard the decline in rootability associated with aging (Libby et a/. 1972). Cuttings collected from the pruned crowns of adult Douglas-fir trees rooted better than shoots collected from unpruned areas of the same trees (Black 1972). The age of trees at first pruning was not reported be­ cause the trees had not been pruned as part of a formal study. They had been pruned at unknown dates to keep branches away from overhead utility lines. Rein­ vigoration likely occurred as a result of the cultural treatments rather than rejuvenation. Another method used to maintain high rooting poten­ tial over a period of years is to serially propagate cut­ tings from other rooted cuttings. Serially propagated adult Douglas-fir clones rooted better (45%) than did shoots taken from the original ortets (I%) or from grafts (9%) (Black 1972). How many years Douglas-fir can be serially propagated and maintain a juvenile or rein­ vigorated condition is unknown, nor whether they can retain such a condition if annual pruning or hedging is delayed I to 4 years after rooting. Although a wealth of information exists on rooting Douglas-fir, information is lacking in several fields or published results are conflicting. This study provides information that will help elucidate the problem areas. Annual rooting trials from 1974 through 1981 were evaluated. The major objectives were to provide infor­ mation on the effects of annual crown pruning on rein­ vigoration of cuttings as demonstrated by increased rooting potential; to determine if rooting potential decreased between ages I 0 and 20; and to evaluate invigoration effects of serial propagation on rooting potential, shoot growth, and growth habit. 10 and 13 years old in 1974 and had been selected for cutting propagation after. graft tests had identified them as being highly graft compatible <Copes 1981 ). Ortet cuttings (CO), consisting entirely of the past season's growth. were gathered in Januury or February from the lower 2 m of each tree. The trees were approximately 4 m tall in 1974 and many grew to greater than 12 m by 1981 . Branch tips and !urge first- and second-order lutcral brunches were pruned for cuttings. From 1974 through 1980, shoots were clipped only from the lower 2 m of each tree; the area above 2 m was allowed unrestricted growth until 1981. In 19!! I shoots were gathered !r' om both the lower 2-m pruned area and the previously unpmned whorl of branches immediately above the 2-m pruned area. Cuttings for serial propagation (CC) were gathered from 220 rooted ramcts of II of the same 14 clones mentioned above. The ramcts had been rooted in 1972 unci 1973 and planted in the field in 1975 and 1976. They were not pruned until 1979. At that time they were about 2 m high. Cuttings were pruned annually after 1979. Shoots were stored in plastic bags at 1- zoe for 1-30 days after field collection. Standard rooting procedure was as fol­ lows: trim the cuttings to 10- to 15-cm lengths. submerge them in a satmated solution of Captan fungicide for I min, dip the basal 2-3 em of each cutting in 0.5% indolebutyric acid (5000 ppm) in 50% ethanol for 5-20 s. and insert the lower 2-4 em of each cutting in the rooting medium. Needles were not removed from the portion of stem buried in the rooting medium. No hormone was used before 1975. Cuttings were reared in two plastic covered houses located outdoors. Cuttings were rooted in two 1.8- by 16.5- by 0.7-m raised beds. Temperature at the cutting base was maintained at 21 ± 3°C with lead heat cables. Watering and humidity were controlled automatically with Mist-A-Matie1 nozzles. A fine grade of mason's sand was used us the rooting medium in 1974 and 1975; a 2: I mix of sphagnum- peat moss and mason's sand each year after 1975 (Copes 1977). Weekly application of dilute concentrations of one of four fungicides (Caplan. Fcrbam, Bcnlate, and Banrot) were used to control root rot and foliage diseases. A complete, water­ soluble fertilizer (20-20-20) was applied weekly after roots were first visible on the base of the cuttings. Cuttings re­ mained in the propagation beds until November m· January, when the rooted cuttings were lifted and transplanted into outdoor beds for I additional year of growth. Yearly observations on rooting success were made in November or January. Analysis of variance tests were used to evaluate annual rooting potential, to compare percentage rooting of CC with annual pruned CO from 1979 through 1981, to compare 1980 shoot growth rate and stem angle of CC and CO cuttings after one season in the transplant beds, and to evaluate the reinvigoration effects of annual pruning on rooting potential. All percentage data in Tables I, 2, and 3 were converted to arc sine values before analysis. Growth and stem angle were measured in September 1980 on CC and CO cuttings, which had grown one season in the transplant beds. Methods 2Trade or proprietary names are included for information purposes only and do not imply any endorsement by the Canadian Journal of Forest Research and the United States Department of Agriculture. Annual rooting trials with 14 coastal Douglas-fir clones were started in February 1974 and were continued with the same clones through 1981 at Corvallis, Oregon. Ortcts were COPES I . Average annual rooting percentage and, in parentheses, number o f cuttings tested f or the years 1974 through 1981 TABLE Clone No. 3 4 14 20 23 25 97 102 106 109 172 197 223 241 X/"i.N " 1978 TABLE 421 Year of rooting" 1975 1974 5 1(154) 41(308) 57(336) 36(1401) 78(96) 80( 104) 33(235) 66(123) 17(416) 83(245) 40(319) 41(235) 16(248) 23(349) 73(2 10) 18( 186) 60(216) 27( 196) 49(252) 59(188) 32( 155) 58(48) 45(284) 88( 172) 49(255) 61(163) 16(134) 37( 134) 49(486) 37(459) 15(407) 27(541) 58(585) 38(50 1) 49(472) 32(219) 42(4569) 44(2593) X 1979 1977 1976 1980 61(7,50) 55(720) 65(1005) 67(630) 58(360) 76(693) 37(515) 62(1080) 68(600) 76(2500) 64(630) 7 1( 1190) 59(690) 66(660) 64(450) 58(255) 65(562) 62(539) 52(360) 65(205) 32(559) 56(660) 77(554) 72(720) 38(419) 23(440) 74(660) 74(406) 66(406) 25(406) 66(406) 49(406) 62(406) 41(406) 66(406) 42(406) 74(406) 67(406) 36(406) 31(406) 62(406) 49(6463) 54(5684) 66( 12023) "i.N 1981 71(1035) 68(570) 68(360) 46(300) 65(360) 73(876) 70(954) 56(864) 63(612) 62(856) 81(864) 36(571) 77(864) 65(864) 82(864) 67(876) 74(852) 34(864) 82(864) 64(3332) 55(3288) 52(3796) 55(4325) 57(2915) 67(2961) 38(2913) 63(3400) 52(2570) 77(5776) 64(3776) 61(3625) 37(3082) 66(3433) 60(6215) 67(11645) 59(49192) data nre not pre cnted bccnuse of excessive loss f'rom disease. 2. Percentage rooting potentials of II clones for cuttings from ortets (CO) and cuttings from rooted cuttings (CC) (number of cuttings in parentheses) Year of rooting Clone No. 3 14 20 23 97 102 106 109 172 197 241 gj"i.N X/"i.N co 72( 173) 74(221) 50(165) 57(180) 22(77) 59(19 1) 8 1(229) 82(303) 59(153) 74(200) 74(193) X cc 49(68) 54(293) 83(168) 60(156) 55(62) 63(385) 29(80) 8 1(256) 69( 183) 59(479) 58(170) 62(2300) 68(2085) 65(4385) co 58(600) 64(407) 62(359) 55(420) 35(4 19) 42(360) 39(360) 77(760) 70(480) 68(360) 56(555) "i.N 198 1 1980 1979 cc co 60(289) 62(360) 50(330) 41(740) 25(140) 45(849) 14(687) 64(987) 55(239) 71(420) 56(622) 59(5080) 49(5663) 54(10743) Each clone was treated as a replicate of the performance of the various treatments. Split plot analysis of variance tests with years as the split plot effect was used on the data in Table 2. In this case clones were used as replication. Rooting data for 1978 were not presented because excessive loss from disease prohibited accurate assessment of rooting. The average loss for the other years was small and fairly constant bewcen years. Results Rooting potential of annually pruned ortets did not decrease when tree age increased from I 0 and 13 years in 1974 to 17 and 20 years in 1981. Instead, rooting 86(420) 67(420) 67(168) 73(420) 31(420) 73(420) 63(420) 83(420) 72(420) 83(420) 86(420) cc 59(420) 46(420) 66(420) 5 1(420) 48(168) 81(420) 66(420) 81(420) 62(420) 84(420) 79(420) 67(4368) 71(4368) 69(8736) co cc 70(1 193) 67(1048) 61(692) 63( 1020) 32(916) 59(97 1) 59(1009) 80(1483) 69(1053) 76(980) 69(1168) 58(777) 53( 1073) 63(918) 46(1316) 40(370) 62( 1654) 33(1187) 7 1(1663) 62(842) 70( 1319) 64(1212) 65(1 1533) 58(1233 1) potential increased from 42 to 67% (Table I). Current­ year rooting exceeded that of the preceding year in five of six comparisons. The increase averaged about 3% each year, although the greatest increase occurred be­ tween 1974 and 1977 (Table I). Annual rooting data for all 14 clones are presented in Table I. Clones and years (linear effect) were both significant at P s:; 0.000 I. Comparisons within and between clones are more accu­ rate for 1979 through 1981 because of greater within­ clone replication. Mean repeatability (r2) for years 1979 to 1981 was 56%, but only 25% for 1974 through 1977. CAN. 422 J. FOR. RES. VOL. 13. 1983 TABLE 3. Comparison of rooting percentages for cuttings TABLE 4. Second-year total height and growth habit (number from pruned and unpruned crown areas of I 0 ortets taken in 198 1 (number of cuttings in parentheses) of degrees the leader deviated from the vertical) for cuttings established from annually pruned ortcts (CO) and from other previously unpruncd rooted cuttings (CC) Type of crown area sampled Ortet No. Pruned Unpruned 4 14 97 102 106 109 172 197 223 24 1 48(430) 67(430) 31(430) 73(430) 63(430) 83(430) 72(430) 83(430) 27(430) 86(430) 22(90) 60(90) 31(90) 8 I (90) 42(45) 88(135) 52(90) 61(90) 2 1(90) 62(45) X/'kN 63(4300) 54(855) In 1979, 1980, and 1981, CC were compared with CO for II of the same 14 clones. Average annual rooting in all 3 years by CO was greater than by CC of the same year (Table 2). Pruning of rooted ramets for serial propagation was not started until 1979, but ortets were pruned annually after I 974. The difference in rooting potential between CO and CC was significant (P :s 0.05). Significance for clones was P ::; 0.000 0 I ; for linear effects, P :s 0.05; and for quadratic year effects, P :5 0.0001. Interactions between type of cut­ ting and years were not significant. In 1981 the effect of annual pruning on rein­ vigoration or maintenance of juvenility was evaluated by comparing rooting behavior of shoots collected from both pruned and unpruned crown areas of the same 10 ortets. Shoots for the unpruned treatment were gathered from branches immediately above the lower 2-m pruned area from which cuttings had been taken each year since 1974. Cuttings from the unpruned areas were larger than those from the pruned areas and apparently more vigorous, but 63% of the shoots from pruned areas rooted whereas only 54% of the shoots from the un­ pruned areas of the same trees rooted (Table 3). The effect of pruning was significant (P :s 0.05). No growth tests after rooting were made on cuttings after rooting. Rooting percentage by cuttings from unpruned areas was greater in only 2 of the 10 ortets. The two clones, 102 and 109, had been among the best rooting clones in 1974 when cuttings were collected from pre­ viously unpruned ortets (Table I). They were not influ­ enced by time or pruning and still rooted readily 7 years later. If data from the two good rooting, nonresponding clones are deleted from the comparison, cuttings from the pruned areas rooted 60 vs. 43% from unpruned areas. This 17% difference more accurately illustrates the increase that can be expected from a majority of the Height (em)" Growth habit n'' Clone No. co cc co cc 3 4 14 20 23 25 97 102 106 109 172 197 223 24 1 39 16 23 22 20 30 28 !9 25 33 32 16 18 25 40 21 27 47 32 46 32 18 32 32 26 36 19 23 44 56 50 47 57 60 50 55 65 45 41 47 46 45 46 57 57 50 49 43 62 52 59 72 60 54 X/'kN 25(350) 29(350) 54( 140) 51(140) 1'Each value is average of·measuremcnts from 25 trees. 1'Euch value is average or measurements from I 0 trees. clones after they have been pruned for several years. Height and stem angle were measured in 1980 on ramets of all 14 clones after I year in the transplant beds. The CC ramets averaged only 4 em taller than the CO ramets (Table 4). This difference was significant statistically (P :5 0.00 I ). Differences between clones and the clone by type of cutting interaction were also significant (P :s 0.001). Difference in annual shoot growth between CC and CO ramets cannot be attributed to the lengths of cutting placed in the rooting beds the previous year because both types had been trimmed the same. Shoots from ramets, however, had larger stem diameters than those from ortets at the time of place­ ment in rooting beds. The CO shoots were only 75% as heavy as those from CC (2.62 vs. 3.54 g/cutting). Measurements of plagiotropism after I year of growth in the transplant beds indicated that cuttings of I I of 13 clones were slightly more orthotropic for CC than for CO. Serially propagated ramets averaged 51 degrees from the vertical, whereas CO ramets averaged 54 degrees (Table 4). The difference between CC and CO was significant (P :s 0.01). The two-way inter­ action between clones and type of cuttings was also significant:(? :5 0.001). Additional evidence of reinvigoration or mainte­ nance of juvenility by annual pruning was seen in 1981 branch morphology observations on ortets, rooted ram­ ets, and grafts. Presence of Jammas growth (2nd tlush branch growth) normally occurs in trees less than I 0 years old, although heavy rainfall in June and July or · COPES heavy application of fertilizer can at times induce older trees to develop lammas shoots (personal observations). Abundant lammas growth in mature trees is unusual. In I981 , lammas growth was found on pruned areas of seven of eleven 17-year-old ortets, whereas none were found on unpruned areas of the 7 trees, or on either pruned or unpruned areas of the other four ortets. Ob­ servations on the rooted ramets used for serial propaga­ tion revealed lammas growth on 2\5 of 220 ramets. All 215 ramets that formed lammas shoots had been pre­ viously pruned, but the 5 ramets that had not formed lammas shoots had never been pruned. Examination of crowns of 6-year-old, unpruned grafts of the same clones revealed no lammas growth. Discussion Juvenile and adolescent Douglas-fir trees can be propagated for rootstock in seed orchards and for com­ mercial Christmas trees. The high rooting potential ex­ hibited by most juvenile and adolescent clones makes this possible. About 67% of over II 000 cuttings were rooted in I 981 when they were 17 and 20 years old. lt is significant that no decrease in rooting potential oc­ curred when tree age increased from I 0 to 20 years. The expected response would have been a decrease. The absence of decreased rooting potential is probably the result of invigoration from annual pruning. Roberts and Moeller ( I 978) found similar results with I 2- to 16-year-old trees. The actual amount of invigoration from annual pruning of ortets is hidden in the 25% increase in root­ ing potential. A portion of the 25% increase resulted from better control of disease and rooting environment during succeeding years of the study. The amount of int1uence from these environmental factors is estimated as the I 2% difference between rooting of shoots from unpruned trees in 1974 and unpruned areas of the same trees in I 98 I (42 vs. 54%). The remaining portion (l3%) is attributed to the invigorating intluence of annual crown pruning. The maintenance of high rooting potential during Douglas-fir adolescent years supports Roberts and Moeller's ( I 978) findings, that rooting potential of 12­ to 16-year-old Douglas-fir trees did not decrease but simply varied annually from 63 to 75%. The 12- to 16-year-old trees had been pruned annually for rooting trials. Quite different results were reported by Black (1972), who worked with unpruned 14- to 20-year-old trees. He found that rooting averaged less than 5%. Much of the difference between Black's results and those of this study and Roberts and Moeller's may have resulted because Black's cuttings came from crown areas that had never been pruned. No increase in rooting potential occurred in unpruned areas. Rooting of cuttings from branches immediately 423 above the pruned zone was significantly less than of cuttings gathered from pruned areas of the same trees. The lack of transfer of reinvigoration factors to unpruned areas, as evidenced by increased rooting po­ tential, supports Black's (1972) findings on partially sheared trees under utility lines. No invigoration effects from serial propagation were detected other than slightly greater shoot and less severe plagiotropism. Cuttings from rooted ramets actually ex­ hibited less rooting potential than CO. One possible reason for the difference was that pruning of ramets was avoided during the 3 or 4 years immediately after rooting. Time for unrestricted growth was needed for the crown of each ramet to enlarge. Ortets were pruned annually during that 3 to 4 years, and consequently, CO continued to increase in rooting potential. The lack of a positive rooting response to serial propagation is con­ tradictory to Roberts and Moeller's (1978) results. They serially propagated six mature Douglas-fir trees and detected positive effects of reinvigoration as evidenced by increased rooting potential. They may have obtained positive results, while I found none because the ortets in my study had been pruned annually. The standard against which they evaluated their serially propagated cuttings was the rooting of cuttings taken from pre­ viously unpruned ortets. A 3-year unrestricted growth period following initial rooting preceded the collection of shoots for serial propagation in both studies. Pruning was usually effective in stopping or slowing maturation. Only the rooting of cuttings from ortet 223 shows a continuous downward trend and, even that clone's serially propagated cuttings, in I981 showed positive signs of increased rooting potential over that of the previous 2 years. Future study will be required to determine if additional time is needed for serial propa­ gation to show a positive effect on average annual rooting. Pruning also promoted or maintained the juvenile growth and vigor that result in the formation of abun­ dant lammas shoots. The relation of juvenility or youth­ ful vigor to presence of lammas growth in Douglas-fir is well known, but the correlation between lammas shoot growth and rooting of stem cuttings in 17- to 20-year-old trees is not clear. ln 1981, several of the ortets with the greatest rooting potential did not form lammas shoots, but several of the poorer rooting ortets did. Thus, the presence of lammas shoots normally denotes juvenile growth or vigor, but lammas does not indicate that such trees have a higher rooting potential than similar trees that do not form lammas shoots. Growth rate and habit of cuttings after rooting is of practical importance because it directly affects how the cuttings can be used. Rooted cuttings that grow slowly or are prone to severe plagiotropism are less desirable than are faster growing, upright trees. This is true for 424 CAN. J. FOR. RES. VOL. 13. 19H3 the production of rooted cuttings for the Christmas tree industry or for compatible rootstocks. Growth in the transplant beds the year after rooting showed good growth for both types of cuttings. The CC were only slightly taller and more upright in growth habit than were CO. The time required for plagiotropic Douglas-fir ramets to become orthotropic is correlated with tree age. Cut­ tings propagated from 3-year-old seedlings all became orthotropic the year after rooting (Franclet 1974), whereas cuttings from 4- to 8-year-old ortets required about 7 years before most ramets became orthotropic (Copes 1976). Field tests will be necessary to determine the number of years required for ramets of 17- to 20-year-old trees to become upright. The required time might be less for CC than for CO because annual pruning appears to have slightly increased the vigor of CC ramets. BHELLA, H. S., and A. N. ROBERTS. 1974. The influence of photoperiod and rooting temperature on rooting of Douglas-fir ( Pseudotsuga menziesii (Mirb.) Franco). J. Am. Soc. Hortic. Sci. 99(6): 551-555. --- 1975. Bud and cambial activity in Douglas-fir as related to stem cutting rootability. For. Sci. 21(3): 269-275. BLACK, D. K. 1972. The influence of shoot origin on the rooting of Douglas-fir stem cuttings. Comb. Proc. Int. Plant Prop. Soc. 22: 142-157. --- 1973. Influences of shoot origin and certain post scverence treatments on the rooting and growth character­ istics of Douglas-fir (PseudOisuga menziesii (Mirb.) Franco) stem cuttings. Ph.D. thesis. Oregon State Univer­ sity. BRIX, H., and H. BARKER. 1971. Rooting of Douglas-fir cuttings by a paired-cutting technique. Proc. Int. Plant Prop. Soc. 17: 118- 120. --- 1973. Rooting studies of Douglas-fir cuttings. Can. For. Serv., Pac. For. Res. Cent. Inf. Rep. BC-X-87. COPES, D. L. 1976. Comparative leader growth of Douglas­ fir graft, cuttings, and seedlings. U.S. For. Serv. Tree Plant. Notes. 27(3): 13- 16. 1977. Influence of rooting media on root structure and rooting percentage of Douglas-fir cuttings. Silvuc Genet. 26(203): 102-106. 1981. Selection and propagation of highly graft­ compatible Douglas-fir rootstocks-a case history. U.S. Dep. Agric. For. Serv .. Res. Note PNW-376. CORNU, D. 1973. Essais prcliminaircs sur Ia selection de clones bouturablcs de Douglas ( Pseudot,\'llg(/ men::iesii Franco). Ann. Sci. For. 30(2): 157-173. FRANCLET, A. 1974. Bouturage du Douglas--enracincment de bouturcs de flechcs prclevccs sur des semis de 3 ans (3 + 0). Rapport Annucl de !'Association Foret­ Cellulosc. pp. 29-43. GRIFFITH, B. G. 1940. Effect of indolebutyric acid, in­ doleacetic acid and alpha napthalcncacctic acid on rooting of cuttings of Douglas-fir and Sitka spruce. J. For. 38: 496-501. HEAMAN, J. C., and J. N. OWENS. 1972. Callus formation and root initiation in stem cuttings of Douglas-fir (Pseudo­ tsuga menziesii (Mirb.) Franco). Can. J. For. Res. 2: 12 1- 134. HYUN, S. K. 1967. Physiological differences among trees with respect to rooting. 14th International Union of Forest­ ry Research Organizations Kongress, Miinchcn. Sect. 22. pp. 168-190. LIBBY, W. J A. G. BROWN. and J. M. FIELDING. 1972. Effects of hedging radiata pine on production, rooting. and early growth of cuttings. N.Z. J. For. Sci. 2: 263-283. McCOULLOCH, W. F. 1943. Field survival of vegetatively propagated Douglas-fir. J. For. 41: 211-212. ROBERTS, A. N., and L. H. FUCI-IIGAMI. 1973. Seasonal changes in auxcn effect on rooting of Douglas-t1r stem cuttings as related to bud activity. Physiol. Plnnt. 28(2): 2 15-221. ROBERTS, A. N., and F. W. MOELLER. 1978. Speculations relating to the loss of rooting potential with aging of Douglns-fir stock plants. Plant Prop. 24( I): 2-4. ROBERTS. A. N., B. J. TOMASOVIC, and L. H. FUCHIGAMI. 1974. Intensity of bud dormancy in Douglns-fir and its relation to scale removal and rooting ability. Physiol. Plant. 31:21 1-216. Ross, S. D. 1975. Production. propagation, and shoot elon­ gation of cuttings from sheared !-year-old Douglas-fir seedlings. For. Sci. 21(3): 298-300. --- --- . •