Tree paeony ( Paeonia suffruticosa ) is one of the most of import cosmetic workss grown in many parts of the universe. The phenomenon of consecutive secondary blossoming ( autumn-blooming ) is normally observed in many horticultural workss including tree paeony. Changing degrees of endocrines and sugars are thought to act upon autumn-blooming tree paeony. Therefore, the aim of this survey was to measure the quantitative alterations in the endogenous endocrines and sugars degrees in the bud samples of ‘ Ao Shuang ‘ cultivar of tree paeony during the flowering period of fall in comparing with that of spring ( 2010 ) . Endogenous endocrines ( IAA, ABA and GA3 ) and saccharides ( sucrose, cut downing sugars and amylum ) degrees were determined for both seasons.
Both endocrine and saccharide degrees showed important different forms in spring season workss from that in fall season workss. Sucrose had the highest concentration in both seasons followed by cut downing sugars and amylum. Increase in sucrose and cut downing sugar contents in fall season workss was accompanied by lessening in starch concentration in fall season workss. Whereas, similar fluctuating forms was observed for those of spring season workss. High concentrations of IAA, GA3 and low contents of ABA were observed in fall season tree paeony and may hold a positive consequence on the blossoming procedure. Therefore, the consequences suggest that quantitative alterations in endogenous endocrines and saccharides may hold a direct influence on the blooming mechanism of fall blooming tree paeony.
Introduction
The tree paeony ( Paeonia suffruticosa ) , native to China, and one of the most brilliant, most beautiful, and most interesting of all workss ( Wister, 1995 ) , has come into its ain as an of import garden works in many states of Asia, America, Europe and Australia.
It is an first-class ornamental works that has been playing an of import societal, economic and medicative function in the lives of the Chinese people. Respected as “ the King of flowers ” in China with a particular cultural symbol of peace and felicity, prosperity and development, power and wealth over many other common workss, the tree paeony has been extensively cultivated in China. Although holding abundant species and cultivars of tree paeony, China is rare in consecutive secondary blossoming cultivars that flower twice or more in the same twelvemonth.
If these rare resources are utilized by coercing them to re-flower in fall will convey benefits in the flower gardening industry. The development of blooming commanding mechanism in tree paeony cultivation is really of import because the works turning in the field flowers merely one time a twelvemonth with short blooming period. Due to the high demand and high monetary value value associated with this works, this phenomenon is unwanted for agriculturists. In a production system of tree paeony with at least 3-5 old ages before bring forthing flowers, with a individual bud for both foliages and flower production and where each planting is usually harvested merely one time a twelvemonth, works quality every bit good as flower measure is of overruling importance for economic grounds and optimum schemes should be ensured in order to obtain maximal output. This creates chances to examine into the blooming physiology of the works under cultivation in order to heighten its production.
Understanding the physiological mechanism of bring oning consecutive secondary blossoming in tree paeony will be indispensable in horticultural industry. Several physiological alterations occur in the shoot vertex that specifically commits the apical meristem to bring forth flowers. The developmental alterations that conveying approximately blooming include endogenous factors, such as sugar alterations, cistrons and endocrines, and external factors, such as twenty-four hours length and temperature. Therefore, blossoming, which represents the look of generative stage, frequently depends on specific environmental and internal developmental signal. The passage from juvenile to adult stage frequently proceeds in response to environmental factors. These factors exert inductive consequence by accordingly doing alterations in sugar and endocrine degrees ( Zeevart, 1979 ) . This stage alteration is a critical phase in works reproductive development because it can be affected by alterations in endocrine and sugar degrees accordingly taking to seed and flower abortion and fruit abscission.
Tree paeony of course experiences a period of cool temperature before the buds turn out to bring forth shoots and flowers. Normally, flower bud induction begins in the perennial ‘ crown ‘ in later summer, as the foliages begin to age, so bloom bud development continues till the works enters quiescence ( Byrne and Halevy, 1986 ) . These buds will barely develop farther until a period of cool temperature has been experienced ( Byrne and Halevy, 1986 ; Fulton et al. , 2001 ; Halevy et al. , 2002 ; Kamenetsky et al.
, 2003 ) . Equally long as sufficient period of cold temperatures have been accumulated to interrupt quiescence, so buds turn out to bring forth shoots and flowers during the warmer temperatures of spring ( Kamenetsky et al. , 2003 ) . This phenomenon is different for some paeony cultivar such as ‘ Ao Shuang ‘ which can besides blossom in summer in the absence of cool temperature. Peony, like many other blooming workss requires endocrines for the growing and normal care of their physiological and biochemical procedures. As organic substances that regulate works ‘ s growing, endocrines play a cardinal function in several works developmental procedures and advance a figure of desirable effects such as embryogenesis, sidelong root development, vascular distinction, apical laterality, clime responses, and flower development ( Friml, 2003 ; Katia and Gilberto, 2004 ; Ana et al. , 2004 cited by Liu et al. , 2008 ) .
As a consequence, endocrines are believed to strongly take part in blooming mechanism in most perennial workss including in tree paeony. However, the consequence of these endocrines varies with environmental conditions at different period during the season ( Koshita et al, 1999 ) . For case, harmonizing to Altman and Goren ( 1972 ) , IAA delays summer bud germination, while GA enhances it and ABA wholly suppress it. Of recent clip, cytokinin profiles in different works variety meats have been reported to differ systematically with seasons.
The degree of cytokinin has been reported to be minimal in mid-June and maximal in late summer in apical buds of Abies nordmanniana. Subapical buds showed the same June lower limit but peaked in mid fall at a much lower degree ( Rasmussen et al. , 2009 ) . In add-on, an repressive consequence of exogenic IAA and ABA, and endogenous IAA were reported in Pharbitis nothing by wijayanti et Al ( 1997 ) in which exogenic GA induce blossoming. Besides endocrines, works growing and development are besides influenced by alimentary handiness. Numerous surveies have shown that saccharose, glucose and fruit sugar are the major assimilate required by most workss. Sucrose is the chief saccharide transported in the workss that provides energy and C skeletons needed for the synthesis of compounds like amino acids, lipid and secondary metabolites.
It besides plays a major function in works metamorphosis as an of import storage sugar and the chief signifier of decreased C translocated from foliages ( beginning ) to developing growing Centre ( sink ) in works ( Heldt and Heldt 2005 ) . Blooming is normally associated with sugar alterations. The change in the handiness of soluble sugars, such as sucrose, can assist modulate many different procedures.
Parker ( 1962 ) ; Li et Al ( 1965 ) ; Sauter and Kloth ( 1987 ) ; Nelson and Dickson ( 1981 ) have reported sugars accretion in woody works tissues in response to seasonal fluctuation. For case, maximal sugar content in winter has been reported and as workss deacclimate in the spring, a lessening in sugar content was observed ( Parker 1962 ; Nelson and Dickson, 1981 ; Fege and Brown, 1984 ; Bonicel et al. , 1987 ) . Research on the relationship between hormonal alterations and flowered bud development may be of importance for the preparation of protocol for a year-around blossoming programme in tree paeony.
Although few research have reported on the fall blooming mechanism of tree paeony ( Jiang et al. , 2007 ; Li, 1998 ; Xiao et Al, 2001 ) , alterations in endogenous endocrines has non been documented and no surveies have included a comparing of endogenous endocrines and sugars between spring season and fall season blooming tree paeony species. Therefore, this research was conducted to finding the dynamic alterations in endogenous endocrines and sugar degrees in blooming mechanism between early-flowering tree paeony that flowers after traveling through cold period and autumn-blooming tree paeony that grows on the same field but flowers in the absence of cold period during the twelvemonth. Information from this research will assist in puting up a protocol by which re-blooming ( ‘ out-of-season ‘ engineering ) can be enhanced and therefore programming of the harvest in blooming for specific selling day of the months at different times of the twelvemonth will be promoted.
Materials and methods
Plant stuff
The experiment was conducted at the Beijing Forestry University baby’s room, Jiu Feng during the spring ( 2010 ) in comparing to the same cultivar of course grown in the field in summer of 2010. Eighteen workss of 5 old ages old tree paeony, Ao Shuang curriculum vitae. with similar growing energy were chosen for probe in February 2010 and August 2010 for the spring and summer season, severally.
Treatments
Prior to try aggregation, agronomic patterns such as earthening, fertilisation, lacrimation, pruning of lame subdivisions etc were carried out. Leafs defoliation was manually exercised in late August before dormancy-releasing intervention. The dormancy-releasing agent used was GA3 at 500 ppm applied at one dosage on developed buds utilizing a picture coppice. A unstained clean knife was used to reap sample buds from the workss and so rinsed with distill H2O to minimise surface taint. The gathered bud samples were instantly placed in an ice box and upon taken to the research lab, these samples were instantly dipped in liquid N ( N2 ) and stored at -80OC until works endocrine extraction and analysis.
Sampling for works endocrine analysis
Bud samples were collected before and after leaf defoliation and so after GA3 application. Sample aggregation after GA3 intervention, targeted bud developmental phases I, II, III, IV, V, VI, VII and VIII harmonizing to Cheng et Al. ( 2001 ) .
Hormonal extraction and analysis
The extraction of endogenous endocrines was conducted as described by Chen ( 1991 ) , with little alterations. The measure of endogenous degrees of ABA, GA3 and IAA was determined utilizing fresh tissue ( 0. 5g ) of the bud. The works tissue was swot with antioxidant ( Cu ) and 10 milliliter of 80 % cool methyl alcohol until it becomes homogenate and transferred into a trial tubing.
Small sum of PVP was added into the homogenate, so spin the mixture on shaker for 10 min. and incubated at 4 OC overnight. The supernatant was transferred into 10ml tube the following forenoon and spin at 6000 revolutions per minute for 20 min. The residue was washed and re-extracted twice more with 2 milliliters of cold methyl alcohol for another 12 hours, and centrifuged under the same conditions as mentioned above and discarded the dust. The combine infusions, up on adding 2-3 beads of NH3, were evaporated ( 35-40OC ) to the aqueous stage in a rotary evaporator. The aqueous stage was so dissolved by adding some distilled H2O followed by the separation of the mixture into two equal parts. One portion of the mixture was adjusted to pH 2. 5-3.
0 with 1N HCl and so extracted three times with equal volumes of ethyl ethanoate. The combine ethyl ethanoate fraction was evaporated to dryness. The dry portion was diluted with 1ml of 3 % methyl alcohol and 97 % 0. 1 M HAc for the finding of acidic endocrines such as IAA, GA3 and ABA.
Determination of endocrine
Hormonal finding was carried out by utilizing the HPLC method, Agilent 1100 chromatography, C18 tubing ( 250*4.
6 millimeter ) matrix competition. Mobile stage: 3 % methyl alcohol and 97 % 0. 1 M HAc for IAA, GA3 and ABA finding. Wavelength of different endocrines: IAA = 280 nanometer, ABA = 260 nanometer, GA3 = 210 nanometer, and velocity inundation is 1 ml/min, competition with “ outer-standard method ” and the standard sample used is Sigma production.
Soluble sugar extraction and analysis
Fresh foliage stuff ( 1g ) was grind in 20 milliliter of distil H2O and extracted in H2O bath at 80OC for 30 min. The suspension was centrifuged for 10 min.
at 6000 revolutions per minute. The supernatant and extracted residues were collected for the finding of saccharose and cut downing sugars, and amylum, severally. Reducing sugars was determined calorimetrically utilizing dinitrosalicylic acid ; sucrose and amylum was determined by anthrone reagent method with glucose as a criterion utilizing the colorimetric analysis of anthrone method as modified for finding of non-reducing sugars ( Xue and Xia, 1985 ) . The optical density was determined by spectrophotometer.
Statistical analysis
All the statistical analyses were conducted utilizing Statistical Package for societal Sciences ( SPSS ) . The average values of the assorted targeted endocrines and saccharides were taken and a one-way ANOVA was used to find the important between interventions at P & lt ; 0. 05 among the interventions.
Consequences
Hormonal contents in the buds
A seasonal comparing of GA3 degree in buds is presented in fig.
1. GA3 contents in both spring and fall workss showed a peak each, matching to bud swelling phase and shoot development phase, severally. After that, in each instance, the tendency of alterations in GA3 degrees showed similar form for both season workss with little addition observed at phase 8 for fall season workss. IAA degrees in the buds of spring season workss showed two extremums matching to flick emerging ( present 3 ) and blossoming ( present 8 ) phases, whereas IAA degrees of fall season works showed three extremums matching to bud swelling ( present 1 ) , shoot developing ( phase 4 ) and flower bud blossoming ( present 7 ) phases. Interestingly, the high degrees of IAA in fall season workss occurred at the phases where it is low in the spring season workss. There was a difference in the ABA content between the spring season workss and the fall workss at the initial phases of growing development.
After that there was no seasonal difference in the ABA content between the two workss. The ABA content exhibited a high degree at the initial phases of development in spring workss than in fall workss but dropped aggressively at phase 3 and maintained similar tendency as that of the fall workss throughout the growing phases, with a little addition at growing phase 5 in fall workss.
Carbohydrate contents in the buds
A quantitative difference in the alteration patterns among developmental phases in contents of saccharide were observed between spring and fall season workss ( Fig 4, 5, 6 ) .
Sucrose and cut downing sugar degrees in spring season workss showed initial addition, so autumn as buds sprout ( present 2 ) and shoot emerges ( present 3 ) , rise once more as flower buds starts to develop ( phases 4 & A ; 5 ) , so fall once more as flower buds unfastened with a crisp rise at blossoming. Whereas that of fall season workss showed an initial lessening so bit by bit increase as buds sprout and keep a stable alteration forms throughout the probe period ( Fig. 3 & A ; 4 ) . Furthermore, amylum in spring season buds showed initial lessening so after that showed similar alteration forms as that of saccharose and cut downing sugars of spring season workss ( Fig. 6 ) . For fall season workss, amylum content showed an initial addition so later reduced with the flower of new vegetive growing and diminution until flower bud formation, at which clip amylum reached seasonal depressions ( phases 3, 4 & A ; 5 ) ( Fig 6 ) .
Under both temperature regimes, sucrose proved to be the major sugar followed by cut downing sugars and amylum. There were no consistent differences in alteration forms of saccharose, cut downing sugars and amylum in spring season workss whereas in fall season workss, a consistent differences in alteration form among the phases was observed between saccharose and cut downing sugar on one manus and amylum on the other manus ( Fig. 4, 5, & A ; 6 ) .
Discussion
Plants growing and development is based on the production of cells in the meristems and the resulting elongation of these freshly formed cells ( Clark, 1997 ; Cosgrove, 1997 ) . Plant endocrines are known to impact both cell division and cell elongation.
Therefore, there is a possibility that hormones regulate cell division and cell growing in workss, straight or indirectly ( Zeevart, 1976 ; Friml, 2003 ; Katia and Gilberto, 2004 ) . Our consequence showed a important different form in the concentrations of endogenous ABA at the initial phases of growing ( b4 – phase 3 ) ( fig. 1 ) in spring season workss from that of the fall season workss. After that, ABA concentrations showed similar alteration form at the staying phases in both season workss. It is possible that the different form was due to the opposite temperature governments experienced at the initial phases of growing. At the clip of significance different degree of ABA, temperature for spring season workss was really low ( February-March ) and that for fall season workss was high ( August-September ) . Because different degrees of ABA concentration were observed during the initial phases of bud development, the phases at which morphological constitution of the flower bud occurs, where normal constructions and development of blooming are set up while abortion associated with blooming rate is determined to a larger extent in tree paeony ( Cheng, 2001 ) , it is possible that endogenous ABA might be one of the cardinal factor responsible for bud development and subsequent blossoming in fall season tree paeony.
Meaning that, the low concentration of ABA in fall season tree paeony likely resulted in fall blossoming. The high degree of ABA content at the initial phases of growing in the spring season workss is non surprising as the works buds were hibernating or had merely been released from winter quiescence and quiescence has been reported to be associated with high degree of ABA ( Dunstone, 1988 ; Djilianov et al. , 1994 ; Kim et al. , 1994 ; Yamazaki et al. , 1995 ; 1999a, B, 2002 Bhargava 1997 ) .
This consequence suggests that the biosynthesis/catabolism or import/export of ABA is affected by fall season status particularly at the early phases ( August-September ) of bud development of tree paeony. Our consequences are consistent with the consequences by Altman and Goren ( 1972 ) which suggested that ABA wholly suppress summer bud shooting but are inconsistent with the consequences of Southwick and Davenport ( 1987b ) which concluded that endogenous ABA might non play a major function in flower bud formation that was controlled by drouth and cooling interventions. The repressive consequence of ABA in the fall season tree paeony besides agrees with Wijiayanti et Al ( 1997 ) and Marumo et Al ( 1990 ) . However, Harada et Al ( 1971 ) and Nakayama and Hashimoto ( 1973 ) reported a promotive consequence of ABA on blossoming. In this survey, high contents of IAA was observed at two extremums synchronising with leaf outgrowth ( present 3 ) and blossoming ( present 8 ) phases in spring season workss whereas, three extremums co-occuring with bud swelling ( present 1 ) , leaf developing ( phase 4 ) and flower bud gap ( present 7 ) phases was observed in fall season workss ( Fig. 2 ) . Because the fall season workss showed three extremums of IAA co-occuring with phases at which the spring season workss showed a relatively low degree of IAA, it is possible that endogenous IAA might hold been influenced by the fall temperature status. This consequence showed there may be a correlativity between endogenous IAA and flower bud formation, bud branch, and flower bud blossoming of tree paeony and acted on flower bud development and its subsequent blossoming.
The important addition in endogenous IAA degree during bud branch has besides been suggested in other species of workss ( Pilate et al. , 1989 ; Gocal et al. , 1991 ; Koshita and Takahara, 2004 ; Liu et al. , 2008 ) .
IAA proved to be the major growth-promotion endocrine in the fall season workss since it showed high degrees at phases where cell division and cell elongation during works development are active. This suggests that IAA plays the cardinal function of bring oning and advancing cell division and cell elongation in fall season workss. The high content of IAA at these phases is non surprising as it is known for its high alimentary attractive force ability ( Koutinas et al. , 2010 ) and that foods are extremely needed for developmental, physiological and metabolic procedures in workss ( Hartmann et al. , 1966 ; Priestly, 1977 ; Golomp and Goldschmidt, 1981 ) . This is of import since the early, fast development of the leaf aboriginal and of immature foliages during the early growing phase, is a requirement of flower bud induction. For the opposite number spring season workss, IAA extremums were merely shown at leaf outgrowth and blooming phases. It is surprising to observe that the alteration forms of IAA degree in spring season workss and fall season workss were face-to-face about throughout the growing phases.
The concentrations of IAA increased in fall season workss at the same phases as a lessening in IAA concentrations in spring season workss was observed. The same alteration forms were observed in the concentrations of IAA and GA3, and IAA and ABA in fall season workss. It means that the raised IAA content was followed by a autumn in the GA3 and ABA contents in the fall season workss ( compare Fig. 1, 2 & A ; 3 ) .
Possibly, there may be an opposite functional relationship between these endocrines and the care of high IAA/GA3 and IAA/ABA has correlativity with fall blossoming for which farther survey is suggested. A important different form in GA3 degrees was observed in buds of fall season workss from that of spring season workss. GA3 contents in buds of spring season workss showed a peak synchronism with bud swelling phase ( present 1 ) , whereas that of fall season workss coincided with complete foliage outgrowth phase ( present 3 ) ( figure 3 ) . After that both maintained the same alteration tendency with fall season workss being the predominant concentrations. Interestingly, the degree of GA3 in the spring season workss was low at the phases of high degree in fall season workss and vice-versa. The high degree of GA3 in fall season workss coincided with the phase at which foliage has wholly emerged and flower buds can be partly or visibly seen in tree paeony, proposing that high degrees of GA3 might hold influence on flower bud formation in fall tree paeony works. Although old surveies have suggested an repressive consequence ( Davenport, 1983, Su et al.
, 1997 ; Koshita et al. , 1999 ; Cao et al. , 2000, 2001 ; An et al. , 2008 ; Prat et al. , 2008 ) or no consequence ( Garner and Armitage, 1996 ) of GA on flower bud formation, our consequences corroborate with the consequences of Luckwill ( 1974, 1980 ) which attributed flower bud formation in apple and pear to the presence of GA and that its absence was the cardinal factor for the deficiency of flower buds in alternately bearing cultivars of these workss.
Surveies by others have besides attributed blossoming to high degree of GA ( Vlahos, 1991 ; Rebers et al. , 1994 ; wijayanti et al. , 1997 ; Nadia et al. , 2006 ; Liu et al. , 2008 ) , which may be attributed to the growth-promotion consequence of GA in stimulating and speed uping cell division and cell elongation ( Hartmann et al. , 1990 ) .
Carbohydrate composing changed in tree paeony bud tissues. A seasonal interchange in the degrees of saccharose, cut downing sugar and amylum between fall season workss and spring season workss was observed. This substituting form was besides observed between saccharose and cut downing sugars, and amylum of fall season workss. The concentrations of these saccharides in fall season workss increased at the same phases as a lessening was observed for that of spring season workss. The concentrations of saccharose and cut downing sugars in bud tissues were high at the initial growing phases in spring season workss than in fall season works but leveled as buds sprout ( present 2 ) and shoot emerges ( present 3 ) and so continued to be reciprocally related to each other throughout the probe period with a pronounced difference at flower gap phase ( present 7 ) ( Fig. 4, 5 & A ; 6 ) . Different temperature governments experienced by both workss may hold influenced the concentrations of these sugars.
The lessening in amylum degrees was besides accompanied by increased degrees of saccharose and cut downing sugars in fall workss, a form that has besides been reported by Ashworth et Al ( 1993 ) ; Mohammadkhani and Heidari ( 2008 ) . The addition in sugar concentrations may be as a consequence of the debasement of amylum ( Fischer and Holl, 1991 ) , proposing that amylum may hold an of import impact on the accretion of sugars in works cells. In spring season workss, a similar fluctuating alteration forms were observed in the concentrations of saccharose, cut downing sugar and amylum.
This means that spring temperature status induced accretion of saccharose, cut downing sugar and amylum among developmental phases in a rise-and-fall form, whereas fall temperature status induced a stable accretion and debasement of saccharose and reduction, and amylum, severally. Though difference in the alteration forms of saccharides between spring and fall season workss was observed, interestingly, the contents of saccharose, cut downing sugar at bud germination and leaf outgrowth phases ( phases 2 and 3 ) and that of amylum at bud shooting phase ( present 2 ) were equal in both season workss ( Fig. 4, 5 & A ; 6 ) .
This suggests that saccharide hydrolysis is a cardinal factor during bud germination and shoot development. The high and low stable substituting degree forms in fall season workss between saccharose and cut downing sugar, and amylum during the growing period as a consequence likely, of autumn temperature governments might be responsible for fall blossoming in tree paeony. Starch content in fall season workss later reduced with the flower of new vegetive growing, presumptively as a consequence of the use of storage saccharide militias to back up shoot and flower bud development and farther declined until flower bud formation at which clip amylum degree reached seasonal lower limit ( Fig 6 ) . Our consequences are inconsistence with Liu et Al ( 2008 ) which concluded that sugar appeared to be irrelevant to blooming of unnatural chestnut. In both season workss, sucrose proved to be the major sugar followed by cut downing sugars and amylum. Our consequences corroborate with Mitthiesen and Stoller ( 1978 ) ; Ashworth et Al ( 1993 ) ; Spencer et Al ( 2001 ) ; Mohammadkhani and Heidari ( 2008 ) . This suggests that saccharose is the chief saccharide transported in the workss that provides energy and C skeletons needed for the synthesis of compounds like amino acids, lipid and secondary metabolites.
Decision
In this survey, different forms in the alterations in the endogenous endocrines ( IAA, GA3 and ABA ) and saccharide ( Sucrose, cut downing sugars and amylum ) degrees among the developmental phases were observed between spring and fall season workss. Endogenous endocrines and sugars have a major influence on the autumn-flowering. Autumn temperature conditions induced the accretion of IAA, GA3, saccharose and cut downing sugar and besides the debasement of ABA and amylum that later induced blooming in fall season workss of tree paeony. The consequences suggest that fall season works ‘ s developmental phases has great influence on the accretion of IAA, GA3, saccharose and cut downing sugars. Furthermore, the quantitative alterations in endogenous endocrines and saccharides may be influenced by seasonal fluctuation that may modulate different blossoming procedures.
Recognition
This work was supported by the Government of Sierra Leone and the Government of PR China through the Chinese Scholarship Council ( CSC ) ( No. 2007694T10 ) .
We greatly acknowledge — — — for her proficient support. Fig. 1. Comparison of ABA content between fall blossoming and spring blossoming tree paeony ( ABA-S: ABA content in spring workss, ABA-A: ABA content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) . Fig. 2.
Comparison of IAA content between fall blossoming and spring blossoming tree paeony ( IAA-S: IAA content in spring workss, IAA-A: IAA content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) . Fig. 3. Comparison of GA3 content between fall blossoming and spring blossoming tree paeony ( GA3-S: GA3 content in spring workss, GA3-A: GA3 content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) . Fig. 4. Comparison of sucrose content between fall blossoming and spring blossoming tree paeony ( GA3-S: GA3 content in spring workss, GA3-A: GA3 content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) . Fig.
5. Comparison of cut downing sugar content between spring blossoming and fall blossoming tree paeony ( GA3-S: GA3 content in spring workss, GA3-A: GA3 content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) . Fig. 6. Comparison of amylum content between spring blossoming and fall blossoming tree paeony ( GA3-S: GA3 content in spring workss, GA3-A: GA3 content in fall workss, b4: before canceling foliages, S1-S8: Phase 1- 8 ) .