Dry Matter Partitioning, Carbohydrate Composition, Protein Reserves, and Fruiting in ‘Autumn Bliss’ Red Raspberry Vary in Response to Pruning Date and Cane Density

in HortScience

In a 2-year experiment (1994 and 1995), plants of primocane-fruiting red raspberry cultivar ‘Autumn Bliss’ grown in a plastic greenhouse were destructively harvested at different growth stages to determine the effect of pruning date and cane density on dry matter distribution, carbohydrate concentration, and soluble protein concentration in different plant parts. Three summer-pruning dates (early, mid, and late July) and four cane densities (8, 16, 24, and 32 canes/m row) were imposed. Relative root biomass decreased from pruning to first flower stage and remained constant thereafter for all pruning dates. Earlier pruning dates corresponded to earlier fruit production, but yield was significantly reduced on later pruning dates and higher cane densities. Sucrose concentration was higher in fine roots than in suberized roots and had a slight decrease during flowering and the beginning of harvest. Soluble protein concentrations did not differ significantly between pruning dates. Reserve carbohydrates in the root system were unaffected by pruning and cane density, and were rapidly used during active vegetative growth, began to recover just after bloom, and were fully recovered at the end of the season. Our experiment suggested that in red raspberry plants grown under poor environmental conditions, current yield is reduced but there is enough carbohydrate accumulation to support next year's growth.

Abstract

In a 2-year experiment (1994 and 1995), plants of primocane-fruiting red raspberry cultivar ‘Autumn Bliss’ grown in a plastic greenhouse were destructively harvested at different growth stages to determine the effect of pruning date and cane density on dry matter distribution, carbohydrate concentration, and soluble protein concentration in different plant parts. Three summer-pruning dates (early, mid, and late July) and four cane densities (8, 16, 24, and 32 canes/m row) were imposed. Relative root biomass decreased from pruning to first flower stage and remained constant thereafter for all pruning dates. Earlier pruning dates corresponded to earlier fruit production, but yield was significantly reduced on later pruning dates and higher cane densities. Sucrose concentration was higher in fine roots than in suberized roots and had a slight decrease during flowering and the beginning of harvest. Soluble protein concentrations did not differ significantly between pruning dates. Reserve carbohydrates in the root system were unaffected by pruning and cane density, and were rapidly used during active vegetative growth, began to recover just after bloom, and were fully recovered at the end of the season. Our experiment suggested that in red raspberry plants grown under poor environmental conditions, current yield is reduced but there is enough carbohydrate accumulation to support next year's growth.

On cultivated red raspberry plants, biennial canes grow on a perennial root system. Usually primocanes (vegetative) and floricanes (reproductive) are present at the same time. This complex system is highly plastic because the various plant parts compete with each other and can compensate for changes (Fernandez and Pritts, 1994; Oliveira et al., 2004; Snyder and Richey, 1930; Waister and Wright, 1989; Whitney, 1982). Early reports on carbohydrate movement in the raspberry plant associate increased carbohydrates in the Fall with phloem breakdown (Brierley and Landon, 1936). Engard (1939) showed that carbohydrates accumulate in the stems and leaves at the beginning of growth, drop rapidly during growth and fruiting, and are again accumulated in the fall. Surplus sugar within the fruiting cane is produced at the time flower buds appear and fruiting laterals stop to elongate (Brierley and Landon, 1936), which allows reserve carbohydrates in the root system to be replenished for the next year's growth (Whitney, 1982). Jennings and Carmichael (1975) showed that raspberry genotypes with lower Winter hardiness have a high proportion of starch in the roots and very little sugars depending on the weather. In cold-hardy genotypes, starch declines and soluble carbohydrates accumulate in winter in response to decreasing temperatures in the fall. The reverse occurs in spring (Palonen, 1999). However, the interaction between carbohydrate accumulation and cultural practices has not been investigated. Also, no reports exist on the seasonal variation of soluble proteins as an important component of red raspberry plant reserves.

The primocane-fruiting red raspberry production, in which the floricanes are removed in the winter, represents an easy system for studying carbohydrate reserves because there is no competition between primocanes and floricanes. In this model, assimilate partitioning is related to leaf phyllotaxy, and translocation can be acroptal or basiptal according to the developmental stage and source-sink relationships (Privé et al., 1994).

Late-season production based on summer pruning of primocane-fruiting cultivars is a low input system that can give high yields with good fruit quality. Pruning date and intensity can be combined to delay the harvest date of a particular cultivar, but the competition between fruiting and vegetative growth under limiting light conditions does not allow commercial fruit production after December in southwest Portugal (Oliveira et al., 1996, 1998). Apparently, summer pruning is detrimental to plant carbohydrate reserves because photosynthetically active leaves are removed and fruiting occurs under limiting light conditions (Oliveira et al., 2004). Canopy management to manipulate light penetration is a routine management practice to raise and stabilize yields and also to improve fruit quality (Oliveira et al., 1999). We recently showed that pruning date significantly reduces yield and that cane density has an optimum between 16 and 24 canes per meter row (Oliveira et al., 2004).

The objective of this research was to determine the pattern of dry matter partitioning and carbohydrate and protein reserve variation through the cropping cycle of red raspberry ‘Autumn Bliss’ using different cane densities and summer pruning dates.

Materials and Methods

The raspberry crop and the production methods are described in detail in Oliveira et al. (2004). The plants of primocane-fruiting cultivar ‘Autumn Bliss’ were established in Mar. 1993 on a sandy soil inside a nonheated greenhouse located at Herdade Experimental da Fataca on the Southwest coast of Portugal (37°N latitude). Rows were 1.6-m apart with a 0.7-m wide “V” trellis to support the canes. Drip irrigation, fertigation, and pest control were done according to commercial practice.

There were three pruning treatments and four cane densities. Primocanes were pruned at ground level on 1 July, 15 July, and 29 July in 1994 and 3 July, 17 July, and 31 July in 1995. Cane densities of 8, 16, 24, and 32 canes per meter row were imposed 1 month after summer pruning when cane height reached ≈0.6 m. These were equivalent to 5, 10, 15, and 20 canes/m2. Primocanes were cut to ground level in January at the end of harvest and new primocanes were allowed to grow, without thinning, during spring to produce fruit.

The experiment was a split-plot design with four replications, in which pruning dates were the main plots and densities the subplots, and was conducted in 2 consecutive years, 1994 and 1995. To assure uniformity, guard rows were established on each side of the greenhouse and a 6.0-m border on both ends. Each individual subplot consisted of 4-m of row. Biometrics and fruit yield data were collected in the central meter of each subplot. Samples for destructive analysis were collected in the remaining plot area. To establish biomass allocation for each plant part, we used the root-sampling area for plant sampling because after a certain period of time, it was not possible to individualize each raspberry plant. Data were subjected to analysis of variance using Statistica for Windows software (StatSoft, Tulsa, Okla.). For regression analysis, growing degree days (GDD) were calculated using a base temperature of 5 °C (Hoover et al., 1989). Seasonal variation in soluble protein was compared by sampling date on a complete randomized design with four replicates.

Whole plant parts were harvested on the three pruning dates, but only two density treatments were chosen each year, 16 and 32 canes per meter row in 1994, and 8 and 24 canes per meter row in 1995. At every harvest, four samples of two plants were collected per each of the six combinations of pruning date × cane density, which included primocanes and leaves. Four random root samples were also taken at each pruning date × cane density treatment using a metal cylinder (15.5-cm diameter and 35.0-cm depth) and separated into fine roots (diameter <2 mm) and suberized roots (diameter >2 mm). Samples were collected on five dates in 1994 and on four dates in 1995 according to each pruning phenologic stage (Table 1). Data from those samples were treated as dependent variables because they were not set at the beginning of the experiment.

Table 1.

Sampling dates for each pruning treatment according to phenologic stage in 1994 and 1995 experiments.

Table 1.

Leaf and primocane fresh weight was determined immediately after sampling. For dry weight, cane, leaf and fruits were oven-dried (Memmert, Schwabach, Germany) at 70 °C until reaching a constant mass. Leaf area per cane was measured using a leaf area meter (Mark2; Delta-T Devices, Cambridge, U.K.). The above- and below-ground biomass was separated and the samples transported in an insulated container and kept cool overnight. Samples were washed and frozen in liquid nitrogen and freeze-dried at −40 °C for 48 h using a lyophilizer (Martin Christ Gefriertrocknungsanlagen GmbH, Epsilon 2–40, Schwabach Germany). Freeze-dried samples were ground into powder with an electric mill (Cyclotec 1093 sample mill; Tecator, Hoganas, Sweden) with a 40-mesh (0.635-mm) screen.

Starch and sugar assay was determined by the iodine potassium–iodide reaction with 32% perchloric acid as extractant. Powdered samples (20 mg) were extracted with 2 mL perchloric acid (32%) for 20 min. Volume was completed to 20 mL and the extract was left to rest for 1 h. From the clear supernatant, 1 mL was combined with 3 mL distilled water and 1 mL iodine solution giving a total solution of 5 mL of solvent. Starch content was evaluated by colorimetry (590 nm) using a spectrophotometer (Pye Unicam SP6–300, Cambridge, England) and calculated from a starch standard curve (adapted from Rudolph et al., 1988).

Reducing and total sugars were determined by the reducing reaction of potassium ferricyanide into ferrocyanide, which develops a yellow color whose absorbance is measured at 420 nm. Samples (100 mg) were extracted with 6 mL distilled water at 100 °C for 20 min and the volume was then completed to 10 mL. For reducing sugar evaluation, 0.25 mL of the extract, 4 mL of 0.2 m of sodium triphosphate buffer at pH 12.4, and 1 mL of ferricyanide solution were heated during 20 min at 100 °C. For total sugars, the same procedure was used, but 0.5 mL of hydrochloric acid (0.3N) was added before heating (method adapted from Buysse and Merckx, 1993; Gaines, 1973; and Wolf and Ellmore, 1975).

Sucrose concentration was calculated by subtracting reducing sugars from total soluble sugars. Final sugar and starch concentrations were expressed as milligrams per gram of dry weight (mg·g−1dry weight).

Soluble protein assay was determined by the Lowry method modified by Bensadoun and Weinstein (1976). For extraction, 250 mg of each powder sample was macerated with sand in a mortar and pestle with 6 mL potassium phosphate buffer 100 mm, pH 8.0, containing 48 μL of 250 mm phenylmethylsulphonylfluoride in a cold chamber (4 °C) for 5 min. This solution was centrifuged (50,000 g, 20 min, 4 °C) and the supernatant desalted in PD-10 columns containing Sephadex G-25 to remove all components with molecular masses below 5 kDa (Scopes, 1982). Two hundred microliters of the extract was precipitated with 50 μL sodium desoxycholate (1%) and 1 mL trichloroacetic acid (TCA). After 10 min, this solution was centrifuged (26,000 g, 5 min). Copper solution (1 mL) and the reagent (0.1 mL) were added to the precipitate and the reaction occurred, in the dark, for the next 3 h. Absorbance was then measured at 750 nm.

Results

Analysis of variance showed that pruning date had no effect on the pattern of plant dry matter accumulation along the cropping cycle, because all pruning dates presented a similar proportion of mass allocated to each plant part. However, cane density affected total plant dry weight with a fast increase early in the season, a peak at ≈1800 GDD in 1994 and 2600 GDD in 1995, and a relatively stabilization thereafter in the low plant-density treatments (Fig. 1). In high plant densities, the initial dry weight increase was faster but with a tendency for decreasing at the end of harvest.

Fig. 1.
Fig. 1.

Total plant dry mass per root sampling area of two cane densities 16 (....) and 32 (—) canes per meter row in 1994 (A) and two cane densities 8 (....) and 24 (—) canes per meter row in 1995 (B). Y = −4E-0.8x3 + 0.0001x2 − 0.0494x + 35.8; r 2 = 0.84; n = 60 for cane density 16 and Y = −8E-08x3 + 0.0003x2 − 0.0787x + 28.7; r 2 = 0.87; n = 60 for cane density 32 in 1994 and Y = −1E-08x3 + 5E-05x2 + 0.0347x + 62.2; r 2 = 0.70; n = 48 for cane density 8 and Y = −1E-08x3 + 5E-05x2 + 0.0128x + 59.4; r 2 = 0.50; n = 48 for cane density 24 in 1995.

Citation: HortScience horts 42, 1; 10.21273/HORTSCI.42.1.77

Dry weight allocated to fine roots decreased in full growth stage and increased thereafter recovering the initial value at end of the cycle in both years (Table 2). Suberized root had a higher mass increase along the cropping cycle, especially during harvest in 1994. In 1995, suberized root mass was always significantly higher than fine root mass (Table 2).

Table 2.

Root dry weight of fine and suberized roots from pruning to end of harvest in 1994 and 1995 experiments.

Table 2.

Starch concentration in the root system was high at the beginning of growth (0–100 GDD), low during growth and flowering (800–1200 GDD), increasing at fruiting (1600 GDD), and similar to the beginning of growth at the end of the season (Figs. 2 and 3). The initial starch concentration in fine and suberized roots was similar for all pruning dates in 1994 and 1995 except for plants pruned on 31 July in 1995 (Figs. 2 and 3). The regression analysis showed that starch utilization and accumulation in 1994 was similar for plants pruned on 1 July and 15 July, but that plants pruned on 29 July accumulated more starch at the end of harvest. The starch utilization pattern by the roots was similar in 1994 and 1995, although at the end of harvest in 1995, starch was lower than at the start of growth because we could follow the starch accumulation pattern until the end of winter.

Fig. 2.
Fig. 2.

Starch concentration variation from pruning to end of harvest in fine roots, suberized roots and primocanes of plants pruned at July 1 (——), July 15(— – —) and July 29 (......) in 1994. Y = 0.0001x2 − 0.248x + 166.4; r 2 = 0.95; n = 40 for July 1 and Y = 0.0001x2 − 0.269x + 179.1; r 2 = 0.93; n = 40 for July 15 and Y = 0.0002x2 − 0.320x + 185.5; r 2 = 0.90; n = 40 for July 29 on fine roots. Y = 6E-05x2 − 0.129x + 90.7; r 2 = 0.80; n = 40 for July 1 and Y = 6E-05x2 − 0.111x + 87.7; r 2 = 0.82; n = 38 for July 15 and Y = 0.0001x2 − 0.206x + 126.1; r 2 = 0.86; n = 40 for July 29 on suberized roots. Y = 0.039x − 16.95; r 2 = 0.84; n = 32 for July 1 and Y = 0.042x − 14.01; r 2 = 0.80; n = 32 for July 15; Y = 0.045x − 18.70; r 2 = 0.91; n = 32 for July 29 on primocanes.

Citation: HortScience horts 42, 1; 10.21273/HORTSCI.42.1.77

Fig. 3.
Fig. 3.

Starch concentration variation from pruning to winter's end in fine, suberized roots and primocanes of plants pruned at July 3 (——), July 17 (— – —) and July 31 (......) in 1995. Y = −4E-08x3 + 0.0002x2 − 0.375x + 209.9; r 2 = 0.93; n = 32 for July 3 and Y = −3E-08x3 + 0.0002x2 − 0.330x + 195.4; r 2 = 0.95; n = 32 for July 17 and Y = −6E-08x3 + 0.0003x2 − 0.455x + 209.4; r 2 = 0.93; n = 32 for July 31 on fine roots. Y = −3E-08x3 + 0.0002x2 − 0.288x + 149.6; r 2 = 0.91; n = 32 for July 3 and Y = −3E-08x3 + 0.0002x2 − 0.304x + 163.0; r 2 = 0.84; n = 32 for July 17 and Y = − 5E-08x3 + 0.0002x2 − 0.359x + 159.0; r 2 = 0.88; n = 40 for July 31 on suberized roots. Y = 0.015x + 20.77; r 2 = 0.70; n = 24 for July 3; Y = 0.017x + 21.19; r 2 = 0.78; n = 24 for July 17; Y = 0.022x + 21.66; r 2 = 0.91; n = 24 for July 31.

Citation: HortScience horts 42, 1; 10.21273/HORTSCI.42.1.77

The pattern of starch concentration variation of the primocanes was rather different from the roots because starch steady increased during the growth cycle (Figs. 2 and 3). Leaf starch concentration variation was similar for all pruning dates and growth phases and varied between 10 and 20 mg·g−1 dry weight. It was only at the end of harvest that plants pruned on 29 July showed significant higher leaf starch levels than plants pruned early (P = 0.001). Cane density treatments had no effect on starch concentration on roots, primocanes, or leaves.

Reducing sugar concentration was lower in suberized roots than in fine roots and fairly constant along the cropping cycle for all pruning dates in 1994 and 1995 (Table 3). Reducing sugars of primocanes reached a peak at the start of harvest in 1 July and 15 July pruning dates but not in late pruned plants. In 1995, the same pattern of variation and the same concentration of reducing sugars were observed. Cane density treatments had no effect on reducing sugar concentration in all plant parts.

Table 3.

Reducing sugars concentration (mg·g−1 dry weight) from pruning to end of harvest in fine roots, suberized roots, and primocanes of plants pruned at three different dates in 1994 (1, 15, and 29 July) and 1995 (3, 17, and 31 July) experiments.

Table 3.

Sucrose variation along the growing cycle followed the starch pattern with a decrease during flowering and start of harvest (data not shown). Sucrose concentration in the primocanes was also lower than in fine roots and had no clear pattern of variation along the growing cycle. All pruning dates had the same sucrose concentration in the leaves. As for starch and reducing sugars, cane density treatments had no effect on sucrose concentrations in 1994 and 1995.

Soluble protein concentrations did not differ significantly between pruning dates in 1994 (Fig. 4). Within the same pruning date, soluble protein reached a minimum at the first flower stage and a maximum at end of harvest for 15 July and 29 July pruning dates, but remained constant for 1 July pruning.

Fig. 4.
Fig. 4.

Soluble protein concentration in fine roots from pruning to end of harvest of plants pruned on July 1 ( ), July 15 () and July 29 () in 1994. Data are the means of each pruning date. Bars ± standard error.

Citation: HortScience horts 42, 1; 10.21273/HORTSCI.42.1.77

Discussion

The primocane-fruiting red raspberries have two different stages of plant development. In the first stage from cane pruning to the onset of flowering (vegetative phase), the meristematic growing points are the primary sinks with starch being mobilized from the root system. In the second stage, from the onset of flowering onward (fruiting phase), two new processes compete for assimilates: fruiting and root carbohydrate storage.

During the vegetative phase in our experiment, dry matter was partitioned and reserves used in a similar way for all pruning dates. In the fruiting phase, late pruning dates slightly increased above-ground dry matter but with a predominance of vegetative growth to fruiting. In the same experiment reported elsewhere (Oliveira et al., 2004), fruit yield per cane decreased linearly with delayed pruning and with increased cane density, but yield per meter row decreased with pruning and was unaffected by cane density. Therefore, decreased yield per cane in higher densities was an evident response to intercane competition. According to the present data, carbohydrate reserves remained at a similar level at the end of the cycle regardless of intraplant competition. Also at the end of the growing cycle, starch concentration was replenished to the initial level. This pattern of variation was already described for carbohydrate reserves in summer-bearing raspberries (Brierley and Landon 1936; Engard 1939).

The number of flowers that develop varies with the environmental conditions and it is sensitive to assimilate supply (Crandall et al., 1974). Flowering buds of late pruned plants grew under lower temperatures and received less radiation, so the rate of assimilate production was expected to fall. If a plant does not have enough carbohydrates to support the conditional growth capacity of all its organs, then the growth of an individual organ will be a function of its ability to compete for available assimilates with other organs (De Jong, 1999). With vegetative and reproductive growth at a minimum, the rate of assimilates delivered to the cane and to the roots should rise, and this may further suppress flower initiation. The whole plant will adjust to the resultant shift in source-sink balance and more assimilates will go to the root system, which explains the higher starch concentration observed in the late pruning root system. Assuming sink strength as the net amount of carbon assimilated during a certain period, we can admit that under growth-limiting conditions (e.g., later pruning dates), the root system sink was stronger than the fruit sink.

The reactions guiding the fixing and distribution of the initial products of photosynthesis between starch and translocable carbohydrates have also to be taken into account. Direct vascular connections determine primary source sink routes. Vascular connections can significantly modify source/sink and sink/sink relationships, e.g., the evident control in the phyllotaxic links between individual leaves and growing organs at the shoot apex. This seems to be the case for red raspberry as showed by Privé et al. (1994). In our experiment, reducing sugars and sucrose concentration were constant in the root system during all growth phases and for all pruning dates, showing no clear modification on translocation processes and indicating a functional phloem.

We consider that there are two alternatives to further improve raspberry yield: 1) increasing photosynthetic rates (biomass production) or 2) increasing harvest index (dry matter distribution). The success will be higher if harvest index is manipulated by modifying carbon partitioning and allocation patterns. Under low-light conditions, fruit production declined in later pruning dates because the plants allocated resources less efficiently to fruit production.

Higher densities had a detrimental effect on cane flower bud development because carbon was allocated to plant tissues that only contributed to higher vegetative biomass. Because there was no effect of cane density on carbohydrate reserves, we may consider that the photoassimilate surplus in high densities (Oliveira et al., 2004) only contributed to higher above-ground dry matter buildup.

Our data do not explain whether plants pruned on the latest date accumulated root reserves at the expense of fruiting. Although fruits were the most affected, sink and root reserves were high on late pruning dates. Plants left unpruned in Summer 1995 to produce a regular crop in July–August yielded three times more than Summer-pruned plants, but reached similar carbohydrate reserves at the onset of rest (Neto, 1997).The hypothesis of Oliveira et al., (2004) that under low light conditions, current yield is reduced but carbohydrates continue to be accumulated in the root system, is now evident, but a more precise experiment must be done to clarify the physiological responses to low light.

Our data also do not explain the leaf contribution to root carbohydrate increase after fruit harvest. Fruit harvest of early pruned plants finished earlier than in late-pruned plants, but net assimilation in the absence of fruit did not contribute to higher root carbohydrate reserves. Further work on leaf photosynthetic efficiency after fruit harvest is needed.

The soluble protein increase at the end of harvest on late pruning dates showed that specific proteins may be accumulated in the raspberry root system for later use. The specific nature of these proteins and the factors that condition protein accumulation should be further investigated.

Summer pruning had important implications on yield of primocane-fruiting red raspberry ‘Autumn Bliss’ but not on carbohydrate reserves as a result of high within-plant compensation. Because carbohydrate reserves are not depleted, Summer pruning of early cultivars can be used for several years with no detrimental effect induced by successive pruning.

Literature Cited

  • BensadounA.WeinsteinD.1976Assay of proteins in the presence of interfering materialsAnal. Biochem.70241250

  • BrierleyW.G.LandonR.H.1936Some evidence relating to the downward movement of photosynthate in fruiting canes of the red raspberryProc. Amer. Soc. Hort. Sci.34377380

    • Search Google Scholar
    • Export Citation
  • BuysseJ.MerckxR.1993An improved colorimetric method to quantify sugar content of plant tissueJ. Expt. Bot.4416271629

  • CrandallP.C.AllmendingerD.F.ChamberlainJ.D.BiderbostK.A.1974Influence of cane number and diameter, irrigation, and carbohydrate reserves on the fruit number of red raspberriesJ. Amer. Soc. Hort. Sci.99524526

    • Search Google Scholar
    • Export Citation
  • DeJongT.M.1999Developmental and environmental control of dry-matter partitioning in peachHortScience3410371040

  • EngardC.J.1939Translocation of carbohydrates in the Cuthbert raspberryBot. Gaz.100439464

  • FernandezG.E.PrittsM.P.1994Growth, carbon acquisition, and source-sink relationships in ‘Titan' red raspberryJ. Amer. Soc. Hort. Sci.11911631168

    • Search Google Scholar
    • Export Citation
  • GainesT.P.1973Automated determination of reducing sugars, total sugars and starch in plant tissue from one weighed sampleJ. Assoc. Off. Anal. Chem.5614191424

    • Search Google Scholar
    • Export Citation
  • HooverE.LubyJ.BedfordD.PrittsM.HansonE.DaleA.DaubenyH.1989Temperature influence on harvest date and cane development of primocane-fruiting red raspberriesActa Hort.262297303

    • Search Google Scholar
    • Export Citation
  • JenningsD.L.CarmichaelE.1975Some physiological changes occurring in overwintering raspberry plants in ScotlandHort. Res.14103108

  • Neto C.B. 1997. Evolução dos hidratos de carbono e proteína solúvel em framboesas remontantes em cultura protegida. Instituto Superior de Agronomia 76 pp.

  • OliveiraP.B.OliveiraC.M.Lopes-da-FonsecaL.MonteiroA.A.1996Off-season production of primocane-fruiting red raspberry using Summer pruning and polyethylene tunnelsHortScience31805807

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.Lopes-da-FonsecaL.MonteiroA.A.1999Summer-pruning intensity affects on off-season production of primocane-fruiting red raspberriesActa Hort.505101105

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.MachadoP.V.Lopes-da-FonsecaL.MonteiroA.A.1998Improving off-season production of primocane-fruiting red raspberry by altering Summer-pruning intensityHortScience333133

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.MonteiroA.A.2004Pruning date and cane density affect primocane development and yield of red raspberry ‘Autumn Bliss'HortScience39520524

    • Search Google Scholar
    • Export Citation
  • PalonenP.1999Relationship of seasonal changes in carbohydrates and cold hardiness in canes and buds of three red raspberry cultivarsJ. Amer. Soc. Hort. Sci.124507513

    • Search Google Scholar
    • Export Citation
  • PrivéJ.P.SullivanJ.A.ProctorJ.T.A.1994Carbon partitioning and translocation in primocane-fruiting red raspberries (Rubus idaeus L.)J. Amer. Soc. Hort. Sci.119604609

    • Search Google Scholar
    • Export Citation
  • RudolphV.MoebesR.WagenknechtK.ZastrowG.1988Schnelle quantitative bestimmung des starkegehalts in erdbeerjungpflanzen nach einer kolorimetrischen methode [Rapid quantitative determination of the starch content of young strawberry plants using a colorimetric method]Gartenbau35150151

    • Search Google Scholar
    • Export Citation
  • ScopesR.K.1982Protein purification—principles and practice.Springer-VerlagN.Y1-39151-185

  • SnyderJ.C.RicheyH.W.1930Carbohydrate composition of protected and unprotected raspberry canesProc. Amer. Soc. Hort. Sci.27146150

  • WaisterP.D.WrightC.J.1989Dry matter partitioning in cane fruitsWrightC.J.Manipulation of fruiting.ButterworthsLondon5161

  • WhitneyG.G.1982The productivity and carbohydrate economy of a developing stand of Rubus idaeus Can. J. Bot.6026972703

  • WolfD.D.EllmoreT.L.1975Automated hydrolysis of nonreducing sugars and fructosans from plant tissueCrop Sci.15775777

If the inline PDF is not rendering correctly, you can download the PDF file here.

Contributor Notes

This work was supported by Fundação para a Ciência e a Tecnologia, Programa Ciência BD/2814/93, Portugal.We acknowledge Adam Dale for critical review of the manuscript.

Research Assistant.

Researcher.

Professor.

Associate Professor.

Professor.

To whom reprint requests should be addressed; e-mail pnbo@mail.telepac.pt.

  • View in gallery

    Total plant dry mass per root sampling area of two cane densities 16 (....) and 32 (—) canes per meter row in 1994 (A) and two cane densities 8 (....) and 24 (—) canes per meter row in 1995 (B). Y = −4E-0.8x3 + 0.0001x2 − 0.0494x + 35.8; r 2 = 0.84; n = 60 for cane density 16 and Y = −8E-08x3 + 0.0003x2 − 0.0787x + 28.7; r 2 = 0.87; n = 60 for cane density 32 in 1994 and Y = −1E-08x3 + 5E-05x2 + 0.0347x + 62.2; r 2 = 0.70; n = 48 for cane density 8 and Y = −1E-08x3 + 5E-05x2 + 0.0128x + 59.4; r 2 = 0.50; n = 48 for cane density 24 in 1995.

  • View in gallery

    Starch concentration variation from pruning to end of harvest in fine roots, suberized roots and primocanes of plants pruned at July 1 (——), July 15(— – —) and July 29 (......) in 1994. Y = 0.0001x2 − 0.248x + 166.4; r 2 = 0.95; n = 40 for July 1 and Y = 0.0001x2 − 0.269x + 179.1; r 2 = 0.93; n = 40 for July 15 and Y = 0.0002x2 − 0.320x + 185.5; r 2 = 0.90; n = 40 for July 29 on fine roots. Y = 6E-05x2 − 0.129x + 90.7; r 2 = 0.80; n = 40 for July 1 and Y = 6E-05x2 − 0.111x + 87.7; r 2 = 0.82; n = 38 for July 15 and Y = 0.0001x2 − 0.206x + 126.1; r 2 = 0.86; n = 40 for July 29 on suberized roots. Y = 0.039x − 16.95; r 2 = 0.84; n = 32 for July 1 and Y = 0.042x − 14.01; r 2 = 0.80; n = 32 for July 15; Y = 0.045x − 18.70; r 2 = 0.91; n = 32 for July 29 on primocanes.

  • View in gallery

    Starch concentration variation from pruning to winter's end in fine, suberized roots and primocanes of plants pruned at July 3 (——), July 17 (— – —) and July 31 (......) in 1995. Y = −4E-08x3 + 0.0002x2 − 0.375x + 209.9; r 2 = 0.93; n = 32 for July 3 and Y = −3E-08x3 + 0.0002x2 − 0.330x + 195.4; r 2 = 0.95; n = 32 for July 17 and Y = −6E-08x3 + 0.0003x2 − 0.455x + 209.4; r 2 = 0.93; n = 32 for July 31 on fine roots. Y = −3E-08x3 + 0.0002x2 − 0.288x + 149.6; r 2 = 0.91; n = 32 for July 3 and Y = −3E-08x3 + 0.0002x2 − 0.304x + 163.0; r 2 = 0.84; n = 32 for July 17 and Y = − 5E-08x3 + 0.0002x2 − 0.359x + 159.0; r 2 = 0.88; n = 40 for July 31 on suberized roots. Y = 0.015x + 20.77; r 2 = 0.70; n = 24 for July 3; Y = 0.017x + 21.19; r 2 = 0.78; n = 24 for July 17; Y = 0.022x + 21.66; r 2 = 0.91; n = 24 for July 31.

  • View in gallery

    Soluble protein concentration in fine roots from pruning to end of harvest of plants pruned on July 1 ( ), July 15 () and July 29 () in 1994. Data are the means of each pruning date. Bars ± standard error.

  • BensadounA.WeinsteinD.1976Assay of proteins in the presence of interfering materialsAnal. Biochem.70241250

  • BrierleyW.G.LandonR.H.1936Some evidence relating to the downward movement of photosynthate in fruiting canes of the red raspberryProc. Amer. Soc. Hort. Sci.34377380

    • Search Google Scholar
    • Export Citation
  • BuysseJ.MerckxR.1993An improved colorimetric method to quantify sugar content of plant tissueJ. Expt. Bot.4416271629

  • CrandallP.C.AllmendingerD.F.ChamberlainJ.D.BiderbostK.A.1974Influence of cane number and diameter, irrigation, and carbohydrate reserves on the fruit number of red raspberriesJ. Amer. Soc. Hort. Sci.99524526

    • Search Google Scholar
    • Export Citation
  • DeJongT.M.1999Developmental and environmental control of dry-matter partitioning in peachHortScience3410371040

  • EngardC.J.1939Translocation of carbohydrates in the Cuthbert raspberryBot. Gaz.100439464

  • FernandezG.E.PrittsM.P.1994Growth, carbon acquisition, and source-sink relationships in ‘Titan' red raspberryJ. Amer. Soc. Hort. Sci.11911631168

    • Search Google Scholar
    • Export Citation
  • GainesT.P.1973Automated determination of reducing sugars, total sugars and starch in plant tissue from one weighed sampleJ. Assoc. Off. Anal. Chem.5614191424

    • Search Google Scholar
    • Export Citation
  • HooverE.LubyJ.BedfordD.PrittsM.HansonE.DaleA.DaubenyH.1989Temperature influence on harvest date and cane development of primocane-fruiting red raspberriesActa Hort.262297303

    • Search Google Scholar
    • Export Citation
  • JenningsD.L.CarmichaelE.1975Some physiological changes occurring in overwintering raspberry plants in ScotlandHort. Res.14103108

  • Neto C.B. 1997. Evolução dos hidratos de carbono e proteína solúvel em framboesas remontantes em cultura protegida. Instituto Superior de Agronomia 76 pp.

  • OliveiraP.B.OliveiraC.M.Lopes-da-FonsecaL.MonteiroA.A.1996Off-season production of primocane-fruiting red raspberry using Summer pruning and polyethylene tunnelsHortScience31805807

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.Lopes-da-FonsecaL.MonteiroA.A.1999Summer-pruning intensity affects on off-season production of primocane-fruiting red raspberriesActa Hort.505101105

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.MachadoP.V.Lopes-da-FonsecaL.MonteiroA.A.1998Improving off-season production of primocane-fruiting red raspberry by altering Summer-pruning intensityHortScience333133

    • Search Google Scholar
    • Export Citation
  • OliveiraP.B.OliveiraC.M.MonteiroA.A.2004Pruning date and cane density affect primocane development and yield of red raspberry ‘Autumn Bliss'HortScience39520524

    • Search Google Scholar
    • Export Citation
  • PalonenP.1999Relationship of seasonal changes in carbohydrates and cold hardiness in canes and buds of three red raspberry cultivarsJ. Amer. Soc. Hort. Sci.124507513

    • Search Google Scholar
    • Export Citation
  • PrivéJ.P.SullivanJ.A.ProctorJ.T.A.1994Carbon partitioning and translocation in primocane-fruiting red raspberries (Rubus idaeus L.)J. Amer. Soc. Hort. Sci.119604609

    • Search Google Scholar
    • Export Citation
  • RudolphV.MoebesR.WagenknechtK.ZastrowG.1988Schnelle quantitative bestimmung des starkegehalts in erdbeerjungpflanzen nach einer kolorimetrischen methode [Rapid quantitative determination of the starch content of young strawberry plants using a colorimetric method]Gartenbau35150151

    • Search Google Scholar
    • Export Citation
  • ScopesR.K.1982Protein purification—principles and practice.Springer-VerlagN.Y1-39151-185

  • SnyderJ.C.RicheyH.W.1930Carbohydrate composition of protected and unprotected raspberry canesProc. Amer. Soc. Hort. Sci.27146150

  • WaisterP.D.WrightC.J.1989Dry matter partitioning in cane fruitsWrightC.J.Manipulation of fruiting.ButterworthsLondon5161

  • WhitneyG.G.1982The productivity and carbohydrate economy of a developing stand of Rubus idaeus Can. J. Bot.6026972703

  • WolfD.D.EllmoreT.L.1975Automated hydrolysis of nonreducing sugars and fructosans from plant tissueCrop Sci.15775777

All Time Past Year Past 30 Days
Abstract Views 0 0 0
Full Text Views 167 167 4
PDF Downloads 35 35 4