Opportunities for regulation of sugar beet storage root growth

The percentage of sucrose in sugar beet storage root fresh and dry matter is closely related to root structure. It has been suggested that the sucrose content might be increased by using plant growth regulators to modify storage root structure through control of cambial development, cell division and cell expansion. During storage root development correlations were found between the changing phytohormone profiles and the formation of secondary cambia and their subsequent cell division and expansion. Sugar beet root derived cell suspension cultures were used for detailed studies of the roles of endogenous phytohormones. The gibberellin synthesis inhibitor paclobutrazol was tested in cell cultures and whole plants. The observations provide a basis for development of plant growth regulator regimes to optimise sucrose yield from sugar beet.     

The growth and development of the storage root of sugar beet

A study was made of the growth of the storage root of sugar beet as a sugar accumulating organ. The storage root grew by simultaneous cell multiplication and expansion from a series of peripheral secondary meristems laid down during the early stages of development. The weight of water and of non-sugar dry matter per cell increased in proportion to the increase in cell volume. The amount of sugar per cell was proportional to cell volume only during the initial stage of cell expansion up to volumes of about 15 times 10^-8 cm³; thereafter it was less proportional. Thus, average cell size is a major determinant of the sugar concentration of the storage root. The implications of this are discussed.     

Sucrose uptake by sugar beet tap root tissue

Sucrose uptake by discs of mature sugar beet root tissue incubated in [¹⁴C]-sucrose exhibited nonsaturating kinetics over the concentration range of 1 to 500 millimolar. Uptake was inhibited by dinitrophenol, sodium cyanide, low O₂, and penetrating sulfhydryl inhibitors. ATPase inhibitors, sodium fluoride, and oligomycin reduced uptake by 20 and 40%, respectively. Uptake as asymmetrically labeled sucrose ([¹⁴C]glucose) occurred with approximately 80% retention of asymmetry, indicating a nonhydrolytic pathway. Uptake was against a concentration gradient and required metabolic energy.     

Effect of irrigation frequency on root water uptake in sugar beet

A 2-year trial was performed on autumn-sown sugar beet grown in pots in order to study the influence of irrigation frequency on the water used by plants along the soil profile. The outdoor pots, containing one plant each, were 1.3 m high and had circular openings, through which Time Domain Reflectometry (TDR) apparatus wave guides could be inserted. Three irrigation intervals were compared and plants were watered whenever the soil layer explored by roots had lost 30% (SWD₁), 50% (SWD₂) and 70% (SWD₃) of the total available water (TAW). During the irrigation season, the water extracted by the plants from each layer along the soil profile (RWU) was determined by monitoring volumetric soil moisture content (), by TDR. At harvest time, root length density (RLD) along the soil profile was assessed using the Tennant method. The applied irrigation frequencies significantly affected the RWU. With the SWD₃ protocol, irrigation was at longer irrigation intervals (9 days) and watering volumes were as high as 84 mm. In this treatment, the plants lost almost 60% of total water from the lower soil layer (0.6–1.0 m). In treatment SWD₁, the irrigation interval was very short (3 days), and water extraction from 0.0–0.6 m soil depth was 92.0%. In the intermediate treatment, the irrigation interval was 5.5 days and a more uniform water depletion was observed along the root zone, approximately equal between the 0–0.6 and 0.6–1.0 m soil layer. Water extraction of sugar beet plants at the deeper soil layers in response to long irrigation intervals was related to an increase in water uptake efficiency of the deeper younger roots and not to an increase in root length density, which, on the contrary, decreased. This morpho-physiological acclimatization to progressive soil water deficit was coupled with an increase of the root/shoot ratio.     

Adenylate patterns of autumn-sown sugar beet differ from spring-sown sugar beet. Implications for root quality

Sugar beet ( Beta vulgaris L) is generally cultivated using two different planting and harvest patterns. In northern zones, spring sugar beet is sown in spring and harvested in autumn, whereas in subtropical latitudes, autumn sugar beet is sown in autumn and harvested in summer. The industrial quality of the root is frequently higher in spring-sown sugar beet crops. In order to explore physiological changes associated with this fact, this study has been focused on the seasonal changes of adenosine 5'-triphosphate and adenosine 5'-diphosphate levels in the storage roots of sugar beet plants, as an index of its metabolic status. The results obtained correspond to a different metabolic status of spring and autumn sugar beet at the moment of harvest. The adenylate patterns of autumn beets suggested a functional and active respiratory system. On the contrary, the patterns shown by spring beets corresponded to those we would expect to see in plants becoming dormant. The proline and glucose contents, which decrease the industrial quality of the root, and the respiratory rate measured in autumn-sown sugar beets, were nearly twice those of spring-sown sugar beets. The combination of an active respiratory system, which allows the carbohydrate catabolism and the synthesis of stress molecules, with the environmental factors at the time of the harvest, could be the underlying physiological mechanism causing some of the differences between spring- and autumn-sown sugar beet crops.     

黄土高原甜菜叶片生长特性及其对块根产量、含糖量的影响

通过对旱地甜菜叶片生长特性及摘除不同叶组对块根产量,含糖量,显微结构的影响研究,结果表明：甜菜第10-20片叶的叶龄最长,积温最高,是甜菜的主要功能叶;甜菜从第20片叶期起进入块根,糖份增长期,从第55叶期起进入糖份积累期;摘除不同叶组的叶片对甜菜块根产量,含糖量及显微结构均有不同程度降低作用,摘除前期叶组对甜菜块根产量,产糖量,根径减幅较大,摘除后期叶组对块根含糖量,维管束环数,维管束环密度减幅较大;摘除第1-30片叶对甜菜影响最大。     

Chromatin- and nuclei-directed ribonucleic Acid synthesis in sugar beet root

The synthesis of RNA by chromatin-bound RNA polymerase prepared from sugar beet (Beta vulgaris) root tissue is completely dependent on the presence of a divalent metal (Mg²⁺ or Mn²⁺) and the presence of four ribonucleoside triphosphates. Accumulation of labeled acid-insoluble product is inhibited by the addition of RNase and actinomycin D to the reaction. When beet root slices are washed for 25 hours, chromatin-associated RNA polymerase activity increases 7-fold over that of unwashed tissue. This enzyme activity declines with further washing. DNA template availability, as measured by saturating levels of added Escherichia coli RNA polymerase, was also found to follow a pattern similar to that for RNA polymerase. Nearest neighbor frequencies of the RNA synthesized by chromatin isolated from unwashed and washed tissue are different.     

Radial and axial water transport in the sugar beet storage root

To evaluate the contribution of transcellular, apoplastic and symplastic pathways to water movements, horizontal (axial pathway) and vertical (radial pathway) sugar beet root (Beta vulgaris L.) slices were studied. Volume flows (Jv) were measured under hydrostatic and/or osmotic gradients, using a computer-based data-acquisition system. When tissues were tested under hydrostatic gradients (0.3 MPa m^-1) a much more important permeability was observed in the axial pathway, as compared with the radial one. Negative pressure gradients (tensions) were as effective as positive ones in inducing a net water movement. After the establishment of a concentration gradient in the radial pathway (obtained by adding 300 M m^-3 mannitol to the employed solution) an osmotic flux, sensitive to HgCI2, was observed. The inhibitory effect of mercurial compounds was reversed by -mercaptoethanol while [¹⁴C] mannitol unidirectional fluxes were not affected by mercurial agents. In the axial pathway, the presence of a mannitol gradient did not develop a sustained osmotic flux. After an initial Jv in the expected direction, the Jv reversed and moved in the opposite way. It is concluded that, in the sugar beet root, water channels play a significant role in water transfers in the radial pathway. On the other side, water and solutes are transported by a hydrostatic gradient in the xylem vessels. In general, these results extend and adapt to a storage root the 'composite transport model' first proposed by Steudle et al.     

Search for actinomycetes--antagonists of fungi causing sugar beet root rot

Actinomycetes belonging to Streptomyces were isolated from the rhizosphere of sugar beet grown on an infected area after cultivation for many years. 44.1 per cent of them proved to be antagonists of phytopathogenic test fungi. The majority of the antagonists were detected among the Cinereus. The lowest number of the antagonists was detected among the Azureus, the maximum number of the antagonists was observed in the middle and at the end of the sugar beet vegetation period. During various periods of the plant development, the number of the antagonists belonging to different sections changed. The majority of the actinomycetes belonging to Streptomyces were active against Fusarium solani and Botrytis cinerea. The antagonists of the fungi were mainly detected among the representatives of the Cinereus and Helvolo-Flavus.     

An analysis of leaf growth in sugar beet.

The increase of leaf area index (L) was examined in a series of sugar-beet crops grown on different sites (Broom's Barn, Suffolk and Trefloyne, Dyfed) or with different husbandry treatments (sowing dates and nitrogen rates) between 1978 and 1982. The development of L could be described as a function of thermal time using three parameters; D_E, which was essentially an estimate of the thermal time required for crop establishment, and rH_L and D_L, the thermal rate and duration, respectively, of the increase of L. Variations in D_E between seasons and with sowing date were small, but significant; they were attributed to factors affecting the condition of the seedbed. There were much larger variations in rH_L, especially between seasons, sites and crops given different rates of nitrogen fertiliser, and there was a strong negative relationship between rH_L and D_L. Much of the variation in rH_L was associated with differences in the concentrations of nitrogen in the lamina dry matter. Faster rates for rH_L at Trefloyne than at Broom's Barn, and in the crop grown in 1982 as compared with other years, were also partly attributable to particularly warm conditions during the early development of some of the larger, faster-growing leaves within the canopy. The wider application of the relationships established from these experiments was tested with data from a series of crops grown on other sites between 1960 and 1962. The relationships held particularly well for beet grown on soils with high water-holding capacity but not for those on soils of low water-holding capacity.     

An analysis of leaf growth in sugar beet.

The responses of leaf appearance and expansion to temperature in sugar beet were measured under controlled conditions, using ruler and auxanometers, to establish a basis for a subsequent analysis of leaf growth in field crops. The studies showed that leaf appearance rate responded linearly to temperature above 1°C, that leaf expansion rate responded likewise above 3°C and that both rates were zero below these base temperatures. Auxanometer measurements of leaf extension showed that daily rates of expansion of leaf area increased linearly with the daily integral of temperature. However, hourly rates of extension in length alternated with those in width during each 24 h cycle in patterns that were not clearly related to hourly changes of temperature or to the day/night sequence.     

Ethylene is differentially regulated during sugar beet germination and affects early root growth in a dose-dependent manner

Main conclusion

By integrating molecular, biochemical, and physiological data, ethylene biosynthesis in sugar beet was shown to be differentially regulated, affecting root elongation in a concentration-dependent manner. There is a close relation between ethylene production and seedling growth of sugar beet (Beta vulgaris L.), yet the exact function of ethylene during this early developmental stage is still unclear. While ethylene is mostly considered to be a root growth inhibitor, we found that external 1-aminocyclopropane-1-carboxylic acid (ACC) regulates root growth in sugar beet in a concentration-dependent manner: low concentrations stimulate root growth while high concentrations inhibit root growth. These results reveal that ethylene action during root elongation is strongly concentration dependent. Furthermore our detailed study of ethylene biosynthesis kinetics revealed a very strict gene regulation pattern of ACC synthase (ACS) and ACC oxidase (ACO), in which ACS is the rate liming step during sugar beet seedling development.     

Distribution of pectic epitopes in cell walls of the sugar beet root

Immunolabelling techniques with antibodies specific to partially methyl-esterified homogalacturonan (JIM5: unesterified residues flanked by methylesterified residues. JIM7: methyl-esterified residues flanked by unesterified residues), a blockwise de-esterified homogalacturonan (2F4), 1,4-galactan (LM5) and 1,5-arabinan (LM6) were used to map the distribution of pectin motifs in cell walls of sugar beet root (Beta vulgaris). PME and alkali treatments of sections were used in conjunction with JIM5-7 and 2F4. The JIM7 epitope was abundant and equally distributed in all cells. In storage parenchyma, the JIM5 epitope was restricted to some cell junctions and the lining of intercellular spaces while in vascular tissues it occurred at cell junctions in some phloem walls and in xylem derivatives. After secondary wall formation, the JIM5 epitope was restricted to inner cell wall regions between secondary thickenings. The 2F4 epitope was not detected without de-esterification treatment. PME treatments prior to the use of 2F4 indicated that HG at cell corners was not acetylated. The LM5 epitope was mainly present in the cambial zone and when present in storage parenchyma, it was restricted to the wall region closest to the plasma membrane. The LM6 epitope was widely distributed throughout primary walls but was more abundant in bundles than in medullar ray tissue and storage parenchyma. These data show that the occurrence of oligosaccharide motifs of pectic polysaccharides are spatially regulated in sugar beet root cell walls and that the spatial patterns vary between cell types suggesting that structural variants of pectic polymers are involved in the modulation of cell wall properties.     

Single nucleotide polymorphism markers linked to root elongation rate in sugar beet

The aim of this study was to identify single nucleotide polymorphism (SNP) markers genetically linked to root elongation rate (RER) in sugar beet (Beta vulgaris L.). A population of 244 F₃ individuals, obtained from the cross between lines L01 (a low RER) and L18 (a high RER), was phenotyped by measuring RER of 11-d-old seedlings grown in a hydroponic culture. Two DNA bulks of 50 F₃ individuals with extreme phenotypes were used for bulk segregant analysis by restriction-associated DNA sequencing. A total of 20 376 SNPs were identified. Single nucleotide polymorphisms were filtered to reduce the number of the false positive and mapped on candidate chromosomal regions of the B. vulgaris reference genome. One of the total of SNPs selected, SNP10139, was strongly linked to RER (P < 0.01). The pattern of association between the SNP10139 genotype and RER was also evaluated on a breeding line panel comprising 40 low and 40 high RER individuals with different allele frequencies between groups (P < 0.01). The SNP10139 sequence was mapped on the B. vulgaris peptide transporter (PTR) gene, a carrier that influences root elongation in Arabidopsis thaliana. Our results suggest that SNP10139 influence RER in sugar beet, and sequence information can be used in marker-assisted selection programs.     

Persistent effects of seedling treatment on growth of sugar beet in pots

Further evidence is provided that the environment of sugar-beet seedlings, or growth substances applied to seedlings, continues to influence growth when the plants are later in other environments. Sugar-beet seeds were germinated at 20 °C in 8, 16 or 24 h photoperiods of constant light intensity, i.e. with different amounts of total radiation. When the seedlings had two leaves (15–18 days old) they were transferred to large pots in the glasshouse. Some seedlings were treated with (2-chloroethyltrimethylammonium chloride) either sprayed on the leaves or applied to the soil, at different times. The treatments affected areas of individual leaves throughout the growing period; plants raised in 24 h photoperiod had the largest leaves, and those in 8 h photoperiod the smallest. Consequently, 24 h plants had most dry matter and 8 h plants least. Plants given most radiation produced leaves fastest and CCC applied early increased the rate, but as the leaves were smaller, except late in 1967, and died sooner, the leaf area duration was less and so yields were less. CCC applied later did not affect leaf production. There was no interaction between amount of radiation and CCC treatment. Twenty-four hour plants had the greatest net assimilation rate (E) early. CCC decreased E early, but increased it later and more when sprayed on the leaves than when applied to the soil. Some factor, possibly pot size, eventually restricted growth and probably diminished the effect of the treatments applied to the seedlings.     

Effect of plant hormones on sucrose uptake by sugar beet root tissue discs

Abscisic acid (ABA), auxins, cytokinins, gibberellic acid, alone or in combination were tested for their effects on short-term sucrose uptake in sugar beet (Beta vulgaris cv USH-20) roots. The effect of ABA on active sucrose uptake varied from no effect to the more generally observed 1.4-to 3.0-fold stimulation. A racemic mixture of ABA and its trans isomer were more stimulatory than ABA alone. Pretreating and/or simultaneously treating the tissue with K⁺ or IAA prevented the ABA response while cytokinins and gibberellic acid did not. While the variable sensitivities of beet root to ABA may somehow be related to the auxin and alkali cation status of the tissue, tissue sensitivity to ABA was not correlated with ABA uptake, accumulation, or metabolic patterns. In contrast to ABA, indoleacetic acid (IAA) and other auxins strongly inhibited active sucrose uptake in beet roots. Cytokinins enhanced the auxin-induced inhibition of sucrose uptake but ABA and gibberellic acid did not modify or counteract the auxin effect. Trans-zeatin, benzyladenine, kinetin, and gibberellins had no effect on active sucrose uptake. None of the hormones or hormone mixtures tested had any significant effect on passive sucrose uptake. The effects of IAA and ABA on sucrose uptake were detectable within 1 h suggesting a rather close relationship between the physiological activities of IAA and ABA and the operation of the active transport system.     

The effects of temperature on leaf growth of sugar beet varieties

Leaf growth of nine varieties of sugar beet (Beta vulgaris L.) was studied at constant temperatures of 7, 11, 15 and 20·C, using generalised logistic curves fitted to the data to estimate the parameters of growth. The rate of leaf appearance increased linearly with temperature and was the same in all varieties. There were differences between varieties in the weighted mean rates of expansion of leaf area per plant (ā), the temperature coefficient of ā and the leaf area duration (D); these differences were caused more by differences in rates of expansion and final sizes of individual leaves than by differences in rates of leaf production. The growth of the first six leaves produced by each plant was examined in detail. The greater size of successive leaves of plants and genotypic differences between comparable leaves were more attributable to differences in the rate than differences in the duration of leaf expansion. Increasing temperatures increased leaf size because they accelerated the rate of expansion more than they shortened the duration of the expansion phase. It is inferred that all effects arose through differences in the initial sizes of leaves before they unrolled from the shoot apex. Dry matter production was proportional to D but was partitioned more to the storage root at the colder temperatures. This may have been related to the differential effects of temperature on cell division and expansion and the relative contribution of these two processes to the final sizes of the leaves and storage root.