One-Leaf Cuttings as a Model to Study Second Growth in the Potato (Solanum tuberosum) Plant

Second growth is an important physiological disorder of thepotato (Solanum tuberosum L.) plant. A model system to studysecond growth was developed using one-leaf cuttings. Photoperiod,temperature, decapitation and leaf removal treatments were carriedout on the plants from which the cuttings were taken and onthe cuttings themselves. Tuberized, one-leaf cuttings takenfrom moderately-induced plants and exposed to 35 °C afterleaf removal showed 95% second growth within 10 d after treatmentinitiation. Conditions that promoted second growth also reducedstarch and dry-matter content, even in tubers that did not developsecond growth. Cuttings, second growth, potato, Solanum tuberosum L, cv, Bintje, Solanum tuberosum L. cv., Désirée, Solanum tuberosum L. cv., Russet Burbank, tuberization, starch content, dry-matter, heat, photoperiod, decapitation, leaf removal     

Initial Anatomical Changes Associated with Tuber Formation on Single-node Potato (Solanum tuberosum L.) Cuttings

One-node potato (Solanum tuberosum L. cv. ‘Katahdin’)cuttings were used to study early anatomical changes associatedwith tuberization. Starch deposition and the percentage frequencyof cells in mitosis increased in the medullary region of thebud within 1 d after cutting, whereas increases in average cellsize were not detected until 4 d after cutting. Starch depositionand mitosis were the earliest detectable changes in anatomyassociated with tuber initiation. Potato, Solanum tuberosum L., tuber initiation, cuttings, cell enlargement, mitosis, starch deposition     

Tuberization in Potato at High Temperatures: interaction between Shoot and Root Temperatures

Tuber formation in intact potato plants (Solanum tuberosum L.cv. Sebago) was reduced by high shoot or root temperatures andstrongly inhibited when both were high. When both the shootand root temperatures were high, disbudding strongly promotedtuberization. There was a smaller increase with warm roots andcool shoots, but no response with warm shoots and cool roots.When both the shoots and roots were cool, disbudding reducedtuberization. Exogenous GA₃, effectively substituted for thebuds at high temperatures, completely preventing tuberization.In apical cuttings, removal of the terminal bud, but not theroots, reduced the inhibitory effects of high temperatures ontuberization. The experiment indicates that tuber productionmay be controlled by at least three factors: a promoter, whichis not assimilate, produced by the buds at cool temperatures;an inhibitor, derived from the buds, but dependent on warm roottemperatures for its formation; and a second inhibitor derivedfrom the mature leaves and produced in response to warm shoottemperatures. Solanum tuberosumL, potato, tuberization, temperature, disbudding, gibberellic acid     

Far-Red Reversal of Red Light Effect during Long-Night Induction of Potato (Solanum tuberosum L.) Tuberization

The hypothesis that phytochrome is involved in the regulation of potato (Solanum tuberosum L.) tuberization was tested. When 5 minutes of red light were given in the middle of the 16-hour dark period to which whole plants were exposed daily for 14 days before making cuttings, the percentage of tuberization on cuttings decreased. The effect of red light was significantly reversed by 2 minutes of far-red light given immediately after the red in each of two separate experiments. This supports the hypothesis that phytochrome is at least indirectly involved.     

Effects of High Air and Soil Temperature Stress on Growth and Tuberization in Solanum tuberosum

Potato plants (Solanum tuberosum L.) were grown at differentair and soil temperatures to determine the effects of high-temperaturestress on root, tuber, and shoot growth. Cooling the soil (17–27C) at high air temperatures (30–40 C) relieved noneof the visible symptoms of heat stress on shoot growth; norwas the degree of induction to tuberize in leaves increased,as reflected in tuberization of leaf-bud cuttings. Heating thesoil (27–35 C) at cool (17–27 C) air temperatureshad no apparent detrimental effect on shoot growth or inductionof leaves to tuberize. However, in each case hot soil largelyeliminated tuber development. In one experiment stolons grewup out of the hot soil and formed aerial tubers upon reachingthe cool air. When leaf-bud cuttings from induced plants wereused as a model system, high soil temperatures inhibited tuberdevelopment from the buried leaf buds, in the absence of anyroot growth. Apparently the induction of leaves to tuberizeis affected principally by air rather than soil temperature,but expression of the signal to tuberize can be blocked by highsoil temperature. Solanum tuberosum L., potato, temperature stress, soil temperature, tuberization     

Tuberization in Potato at High Temperatures: Responses to Gibberellin and Growth inhibitors

The responses of potato plants (Solanum tuberosum L., cv. Sebago)to high temperatures (32 day/28 C night or 32/18 °C) andgibberellin are similar, in that they promote haulm growth andsuppress tuber production, whereas low temperatures (22/18 °C)abscisic acid and CCC have the opposite effect, promoting tuberproduction and reducing the growth of the haulms. The inhibitoryeffect of the high temperatures on tuber production, under aphotoperiod of 14 h, was almost completely reversed in theseexperiments by the application of CCC, and partly reversed byABA. Single-leaf cuttings from plants grown at the various temperaturesand chemical treatments responded in the same way as the wholeplant. It is suggested that both haulm growth and tuber initiationare influenced by a common hormonal control, and that temperatureexerts its influence by altering the balance between the levelsof endogenous gibberellins and inhibitors. These substancesapparently act directly on the stolon tip, rather than throughtheir general influence on haulm growth. Solanum tuberosum L., potato, tuberization, temperature response, gibberellin, abscisic acid, 2-chloroethyltrimethylammonium chloride (CCC)     

Tuberization in Potato at High Temperatures: Interaction between Temperature and Irradiance

Potato plants (Solanum tuberosum L., cv. Sebago) responded similarlyto high temperatures and low irradiance by diverting dry matterto the shoots rather than the tubers, and changes were notedin a range of morphological characteristics. It is proposedthat the effect of both high temperature and low irradianceis brought about by the increased production of a growth substance,possibly gibberellin, which inhibits tuber formation, and thattuber yield is determined by the balance between temperatureand irradiance. Solanum tuberosum L., potato, tuberization, temperature, irradiance, gibberellin     

Initial Anatomical Changes Associated with Tuber Formation on Single-node Potato (Solanum tuberosum L.) Cuttings: A Re-evaluation

Cell division and cell expansion during early stages of tuberdevelopment were studied using developing axillary buds on single-leafcuttings from potato (Solanum tuberosum L.). Cuttings takenfrom plants induced to form tubers, by short day (SD) treatment,were compared with cuttings from non-induced (long day, LD)plants. In the apical zone of the buds, cell division occurredfrom the first day after cutting, in both LD and SD cuttings.The planes of these divisions were transverse, associated withelongation of the buds. At day 5, a new orientation of celldivision was observed in the subapical zone of SD cuttings only.These divisions were longitudinal, associated with radial growth.Cell expansion occurred in both SD and LD cuttings, and wasnot uniquely related to the onset of tuber formation. Copyright1999 Annals of Botany Company Solanum tuberosum L., potato, tuber formation, cell division, cell expansion.     

Tuberization in Potato at High Temperatures: Promotion by Disbudding

The role of the terminal and axillary buds, as presumptive organsof gibberellin synthesis, in the control of tuberization inpotato (Solanum tuberosum L., cv. Sebago) at high temperatureswas studied. Decapitation alone strongly promoted the outgrowthof axillary buds, but did not promote tubenzation. When growthof the axillary buds was suppressed by the use of chemical pruningagents (MH, TIBA or 1-decanol), tuberization was promoted. Manualremoval of the buds promoted tuberization to a similar extent.The results are consistent with the hypothesis that the budsare major sites of gibberellin synthesis in the potato, andthat high temperatures stimulate the synthesis of gibberellinsand their export to the stolons, where they inhibit tuber formation. Solarium tuberosum L., potato, tuberization, temperature, disbudding, chemical pruning, gibberellins     

Changes in Tuberization and Assimilate Partitioning in Potato (Solanum tuberosum) During the First 18 Days of Photoperiod Treatment

Potato (Solanum tuberosum L.) plants were equilibrated under18-h days (LD) before a subset of the plants was transferredto 10-h photosynthetic periods with either a dark night (SD)or an 8-h dim photoperiod extension with incandescent lamps(DE). Plants were harvested at regular intervals for growthanalysis during the 18 d after transfer. Leaf area increasedrapidly under SD and LD but was inhibited under DE. Internodeelongation was similar under SD and LD, but much higher underDE. Stem d. wts were lowest under SD. Axillary branching wasgenerally greatest under LD. Total shoot weights were greatestunder LD. Total shoot weights were similar under SD to thoseunder DE, even though within 18 d of transfer as much as one-thirdof the biomass of SD plants was in tubers. Tuber initiationwas later under LD than under SD, and was delayed even moreby DE. High temperature increased the delay in tuberizationfrom LD. The early tuber initiation under SD was concurrentwith a rapid increase in leaf area under SD, not with an earlycessation of leaf growth. This was contrary to assumptions basedupon studies of long-term effects of photoperiod. The resultanthigh sink strength under SD contributed to the greater efficiencyof biomass production. Potato, Solanum tuberosum L. cv. Norchip, photoperiod, temperature, morphology, tuberization, growth analysis, biomass partitioning, sink strength, leaf area, short term effects     

Starch Accumulation in Leaves of Potato (Solanum tuberosum L.) during the First 18 Days of Photoperiod Treatment

Potato (Solanum tuberosum L.) plants were grown under long days(LD) of 18 h before a subset of the plants was transferred to10-h photosynthetic periods with either a dark night (SD) oran 8-h dim photoperiod extension with incandescent lamps (DE).Temperature was constant at 21 °C. Leaves were sampled atthe beginning and end of the high density light period for starchanalyses. Potato leaves accumulated starch more rapidly underSD than under LD; and this difference continued after a secondmajor sink, the tuber, began to develop. Starch accumulationover 10 h in SD leaves was three times higher than in LD leaves,even after 17 d of treatment. By this time SD gave higher wholeplant relative growth rates than LD, and the tuber mass of SDplants exceeded 30% of their total plant biomass. The DE treatmentresulted in starch accumulation intermediate to the LD and SDtreatments. Genotypes likewise differed: the earlier genotype,more strongly induced to tuberize, had higher leaf starch accumulationthan the later genotype. The effects of photoperiod and genotypewere also present when potatoes were grown at 27 °C, a temperatureunfavourable for tuberization under LD. Thus the formation ofa strong tuber sink was consistently associated with more rapidleaf starch accumulation. Potato, Solanum tuberosum L., cv. Norchip, photoperiod, temperature, genotype, starch accumulation, partitionin     

Induction and accumulation of major tuber proteins of potato in stems and petioles

A family of immunologically identical glycoproteins with apparent molecular weights of approximately 40,000 are among the major tuber proteins of potato (Solanum tuberosum L.). These proteins, as purified by ion-exchange and affinity chromatography, have been given the trivial name `patatin.' To determine if patatin can be used as a biochemical marker to study the process of tuberization, its amount was measured in a variety of tissues by rocket immunoelectrophoresis and by enzyme-linked immunosorbent assay (ELISA).

Patatin comprises 40 to 45% of the soluble protein in tubers regardless of whether they are formed on underground stolons or from axillary buds of stem cuttings. Under normal conditions, patatin is present in only trace amounts, if at all, in leaves, stems, or roots of plants which are either actively forming tubers or which have been grown under long days to prevent tuberization. However, if tubers and axillary buds are removed, patatin can accumulate in stems and petioles. This accumulation occurred without any obvious tuber-like swelling and would occur even under long days. In all tissues containing large amounts of patatin, the other tuber proteins were also found as well as large amounts of starch.

     

Sucrose application causes hormonal changes associated with potato tuber induction

Stems of potato plants (Solanum tuberosum L. cv. Dianella) were immersed in solutions containing water (control), sucrose, glucose, paclobutrazol, and gibberellic acid. The effects of these treatments on the ethylene release, levels of endogenous gibberellins, glucose, sucrose, and starch were correlated with tuberization of nodal cuttings, excised from potato stems. Paclobutrazol and sucrose improved the percent of tuberization and/or increased tuber weight. In contrast, GA₃ inhibited tuber formation compared with the control. The level of endogenous free GAs was negatively correlated with percent tuberization. However, the level of conjugated GAs was positively correlated with both percent tuberization and tuber weight. The effect of sucrose on potato tuber induction in relation to the possible role of sucrose in GA-conjugate formation is discussed.     

Blue Light Inhibition of Tuberization in a Day-Neutral Potato

In tests on the effects of light quality on potato tuberization, continuous blue light was found to consistently inhibit tuberization of tissue-cultured plantlets of Solanum tuberosum ssp. tuberosum cv. ??Norland??. Other tested cultivars, including sports of ??Norland??, formed tubers under continuous blue light. Microarrays identified BL, GA7ox, and Nudix genes as exhibiting altered expression in response to blue light treatment. Quantitative RT-PCR (qRT-PCR) showed that GA7ox RNA increased in ??Norland?? but not in ??Sangre?? plantlets in blue light compared to darkness. RNA levels of genes identified in the literature as having roles in potato tuberization were also measured using qRT-PCR. Levels of GA20o1x, but not GA2ox, RNA increased in response to blue light in ??Norland?? plantlets. BEL5 RNA content was greater under blue light compared to darkness for both ??Norland?? and ??Sangre?? plants. Levels of FT were not significantly different in blue light compared to dark-treated ??Norland?? plants, but were low in blue light-treated compared to dark-treated ??Sangre?? plants. Addition of ancymidol to ??Norland?? plants exposed to blue light overcame blue light inhibition of tuberization. Ancymidol prevents the oxidation of ent-kaurene to ent-kaurenoic acid, thus inhibiting gibberellin biosynthesis. These data suggest that blue light may increase GA accumulation in ??Norland?? plants, as has been shown to occur in Arabidopsis plants. The novel effect of blue light in inhibiting tuberization of ??Norland?? plants suggests that this system could be a useful tool in further elucidating the mechanisms of day-neutral potato tuberization.     

Tuber Formation and Growth of in vitro Cultivated Transgenic Potato Plants Overproducing Phytochrome B

We studied the effect of the ectopic expression of the Arabidopsis PHYB gene, which encodes the phytochrome B (phyB) apoprotein, under the control of cauliflower mosaic virus 35S promoter on the photoperiodic response of tuberization and growth of potato (Solanum tuberosum L., cv. Désirée) transformed lines. Stem cuttings of transformed and control plants were cultured on Murashige and Skoog nutrient medium containing 5 or 8% sucrose in the phytotron chambers at 20°C under conditions of a long day (16 h), a short day (10 h), or in darkness. We showed that the overexpression of the PHYB gene enhanced the inhibitory effect of the long day on tuberization. In addition, tuber initiation in these transformed plants occurred at a higher sucrose concentration. The insertion of the PHYB gene decreased plant and tuber weights and shortened stems and internodes. Thus, we demonstrated the complex result of the PHYB gene insertion: it affected the photoperiodic response of tuberization, the control of tuber initiation by sucrose, and the growth of potato vegetative organs.     

An Improved Method Involving Hydroponic Culture for the Production of Sexual Hybrids between Dihaploid Solanum tuberosum and Diploid S. microdontum

Dihaploid Solanum tuberosum and diploid S. microdontum plantswere grown in soil and hydroponics under glasshouse and growthroom conditions. A high light intensity was necessary for flowerinduction in both species and the dihaploid flowered only whengrown in hydroponics. Premature berry abscission was retardedby tuber removal and prevented by the addition of indole aceticacid to the nutrient solution. Seeds from prematurely abscisedberries germinated poorly in soil, but germinated almost aswell as those seeds from indole acetic acid-treated plants whenplaced on Murashige and Skoog (1962) based culture medium. Thehybrid plants were intermediate in morphology, compared to theparents, possessed heterotic vigour and were male fertile. Germinatinghybrid seeds on a colchicine-containing medium led to poorlygrowing plants with ploidy chimeras. Hybrid plant ploidy levelswere doubled by regenerating plants from stem/leaf explantson the tuber disc regeneration medium of Jarret et al. (1980). Key words: Solanum tuberosum, S. microdontum, sexual hybridization, hydroponic culture, tissue culture     

Enhancement of Tuberization of Axillary Shoot Buds of Potato (Solarium tuberosum L.) Cultivars Cultured In Vitro

Factors controlling growth and tuberization of axillary budsin shoots of plantlets of potato (Solarium tuberosum L.) culturedin vitro were investigated. Correlative inhibition restrainedgrowth and tuberization of the axillary buds. Exposure of intactplantlets for various periods (4 to 48 h) to low (2 or 12C)or high (30 C) temperatures as comparedto 18C, did not alleviatecorrelative inhibition. Removal of the apical part of the shoot,the roots or both was generally ineffective Elevating sucroseconcentration from 30 to 80 g dm^–3 promoted tuberizationon axillary buds, and the cytokinin 6-(-dimethylallylamino)purine (2iP), alleviated correlative inhibition and enhancedtuberization in intact plantlets. In the whole plantlet mostof the tubers were formed on the basal nodes, however, oncecorrelative inhibition was eliminated by the dissection of theshoot to single node sections, tubers were formed on every axillarybud. The single most effective factor inducing tuberizationin single node sections was the growth retardant ancymidol,an inhibitor of giberellin biosynthesis. Key words: Potato, Solanum tuberosum L., in vitro tuberization, correlative inhibition     

Tuberization in Potato at High Temperatures: Gibberellin Content and Transport from Buds

Warm temperatures (35°C day/30°C night) which inhibittuberization in potato (Solanum tuberosum L., cv. Sebago) increasedgibberellin activity in crude extracts from buds, but not frommature leaves, as determined by the lettuce hypocotyl bioassay.Changes in the growth of tubers and stolons indicate the occurrenceof basipetal movement of GA₃ applied to the terminal bud ora mature leaf. ¹⁴C labelling from GA₃ or mevalonic acid injectedjust below the terminal bud was recovered in the lower shoot,stolons and tubers, but the amount transported was greater atcool temperatures (20/15°C). It is concluded that high temperaturespromote the synthesis of gibberellin in the buds rather thantransport to the stolons. Solanum tuberosum L., potato, tuberization, gibberellin     

Growth and Tuberization of Potato (Solanum tuberosum L.) under Continuous Light

The growth and tuberization of potatoes (Solanum tuberosum L.) maintained for 6 weeks under four different regimes of continuous irradiance were compared to plants given 12 hours light and 12 hours dark. Treatments included: (a) continuous photosynthetic photon flux of 200 micromoles per square meter per second cool-white fluorescent (CWF); (b) continuous 400 micromoles per square meter per second CWF; (c) 12 hours 400 micromoles per square meter per second CWF plus 12 hours dim CWF at 5 micromoles per square meter per second; (d) 12 hours micromoles per square meter per second CWF plus 12 hours dim incandescent (INC) at 5 micromoles per square meter per second and a control treatment of 12 hours light at 400 micromoles per square meter per second CWF and 12 hours dark. The study included five cultivars ranging from early- to late-season types: `Norland,' `Superior,' `Norchip,' `Russet Burbank,' and `Kennebec.' Tuber development progressed well under continuous irradiation at 400 micromoles per square meter per second and under 12 hours irradiance and 12 hours dark, while tuber development was suppressed in all other light treatments. Continuous irradiation at 200 or 400 micromoles per square meter per second resulted in severe stunting and leaf malformation on `Superior' and `Kennebec' plants, but little or no injury and vigorous shoot growth in the other cultivars. No injury or stunting were apparent under 12-dim light or 12-dark treatments. Plants given 12 hours dim INC showed significantly greater stem elongation but less total biomass than plants in other treatments. The continuous light encouraged shoot growth over tuber growth but this trend was overridden by providing a high irradiance level. The variation among cultivars for tolerance to continuous lighting indicates that potato may be a useful species for photoinhibition studies.     

Stomatal Response of some Cultivated and Wild Tuber-bearing Potatoes in Warm Tropics as Influenced by Water Deficits

Leaf resistances of 14 cultivated potato genotypes (Solanumspp) and three tuber-bearing wild Solanum species were comparedwhen plants were grown under water stress at two tropical sitesFactors investigated were diurnal changes in leaf resistance,the effect of plant age, transient drought versus well-wateredconditions of potted and field-grown plants These measurementswere carried out in order to determine the stomatal behaviourof tuber-bearing genotypes and species Significant genotypic differences in leaf resistances were notedwithin the cultivated genotypes All genotypes had higher resistanceswhen water-stressed, but LT-7 appeared to have the lowest leafresistances Genetic differences in stomatal behaviour of tuber-bearingSolanum species were confirmed Abaxial stomatal resistancesof water-stressed plants of the species ranged between 1 74and 13 8 s cm^–1 Stomata of S chacoense were less affectedby drought (three-fold) than S tuberosum (four-fold) The greatesteffect was on S jungasense (five-fold) and on S raphanifoliumThese data show that stomata behaviour among tuber-bearing Solanumspecies is sufficiently different to warrant investigationsof drought-resistance in potato species under dry hot conditions Solanum tuberosum L., Solanum raphanifolium, Solanum chacoense, Solanum jungasense, leaf resistance