Somaclonal variation in tuber traits of potato

Summary About 13,000 somaclones of 17 cultivars and clones of potato were obtained from in vitro callus cultures and individually planted in a greenhouse, followed by tuber generations grown in the field. These plants were subjected to the multistage selection procedure commonly used in potato breeding. Over a period of five years and three field generations the tuber number, size, shape, eye depth, starch content, starch yield and tuber appearance of these somaclones were assessed and compared with that of the controls. These characters varied depending on donor genotype and trait. The frequency of variants was assessed and there were acceptable proportions of desirable abberrants and invariants among the somaclones. Depending on trait the average gain rate for all donor genotypes ranged between 0.2 and 2.3% for-deviants, between 12.2 and 15.5% for invariants and between 0.1–1.4% for +deviants. It is concluded that this technique should be used into potato breeding programmes to improve commercially important characteristics of specific cultivars and breeding clones.     

The effect of early drought stress on numbers of tubers and stolons of potato in controlled and field conditions

Summary Potatoes were grown in plots, pots or containers in a glasshouse or under a rainshelter. Water was withheld from emergence onward or from tuber initiation onward in some treatments. The number of stolons per stem was greatly reduced but the number of tubers + tuber initials per stolon remained unchanged (cv. Radosa) or increased (cv. Bintje) as a result of the earliest drought treatment. The later dry period did not affect numbers of stolons and tubers (+ initials). These findings correspond well with data from field trials which were sampled for 13 consecutive years and which showed a highly significant linear relationship between the number of tubers per stem and total rainfall during the first 40 days after planting, which ranged from 17.5 to 120 mm.     

Observations on the bulking and development of tuber size distribution in maincrop potatoes at Rothamsted in 1964–75

Summary Tuber samples taken at 2-week intervals from plots of cvs King Edward, Majestic and Pentland Crown in 1964–75 showed that the maximum bulking rates ranged from 1 to 5 tonnes/hectare/week. Maximum tuber numbers were usually produced by 11 weeks after planting and thereafter numbers declined. In September, about 70% of the tubers initiated were recovered, and between 24 and 52% of the tubers had reached marketable size (>4 cm). Tuber size distributions developed similarly in all years with all cultivars. During tuber initiation the distributions appeared as one population, and usually between 9 and 11 weeks after planting a second population of larger tubers developed. The population of small tubers later declined although in most years it was still present in September. This development fitted a model based on a mixture of two normal distributions and was determined by five parameters; the proportion of tubers in the small population and the means and standard errors of both populations.     

Some observations of patterns of tuber formation and growth in the potato

Summary Patterns of tuber formation and growth were studied with the varieties Désirée Maris Piper and Peniland Crown. The varieties formed similar numbers of stolons but different numbers of tubers. More tubers were formed at the first node than at any other but Pentland Crown formed fewer tubers over nodes 2–8 than the other varieties. The range of nodes over which tubers formed and competition for photosynthate within a node appeared to be factors controlling the tuber size distribution. The time over which tuber formation occurred was not important. Measurements of individual tuber growth suggested that some tubers followed an approximately sigmoid curve, some grew linearly and some showed periods of tuber growth interrupted by periods when the tubers grew slowly if at all. Estimation of the fresh and dry weights of growing tubers using a relationship between the product of tuber axis lengths and the weights of harvested tubers proved most satisfactory.

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Zusammenfassung In einer Reihe von Versuchen, bei denen die Sorten Désirée, Maris Piper und Pentland Crown verwendet wurden, wurden die Muster der Knollenbildung und das Knollenwachstum untersucht. Diese 3 Sorten haben bei gleichem Pflanzenabstand eine unterschiedliche Verteilung der Knollengr?sse; Pentland Crown neigt zur Produktion grosser Knollen, Maris Piper bringt viele kleinere Knollen und Désirée liegt dazwischen. Um die Knollenbildung zu verfolgen, wurden dreimal in der Woche Proben der wachsenden Pflanzen genommen und die Zahl der Stolonen und Knollen pro Nodium gez?hlt (Tabelle 1). Um das Wachstum der Einzelknollen in Kisten (Fig. 1) und im Feld unter Polythenfolie (Fig. 2) zu verfolgen, wurden die drei Hauptachsen gemessen und daraus das Volumen (in der Annahme, dass die Knolle ein Ellipsoid ist) oder das Gewicht errechnet (wobei das lineare Verh?ltnis zwischen dem Produkt der Knollenachsen und dem Frisch- und Trockengewicht von Knollen verwendet wurde, die unter gleichen Bedingungen gewachsen waren und jede Woche geerntet wurden, wenn die Messungen gemacht wurden). Diese Technik erwies sich als befriedigend und in Tabelle 2 sind die Werte der Parameter für die erste und letzte Messung in jedem Jahr zusammen mit den entsprechenden Bestimmtheitsmassen angegeben. Die Sorten bildeten die gleiche Anzahl von Stolonen aber unterschiedliche Knollenzahlen. Am ersten Knoten wurden mehr Knollen als an irgend einem anderen gebildet, aber Pentland Crown bildete an den Nodien 2–8 weniger Knollen als die anderen Sorten. Das Verh?ltnis der Gesamtzahl der Knollen pro Nodium und der Knollenzahl pro Nodium ausgedrückt in Prozent der Stolonenzahl (Fig. 3) unterschied sich zwischen den Sorten betr?chtlich und ist vielleicht für die unterschiedliche Verteilung der Knollengr?ssen verantwortlich. Die Zeit der Knollenbildung war bei allen Sorten gleich. In den Versuchen über das Knollenwachstum wurden mehr als 500 Knollen gemessen und einige sind in Fig. 4–6 dargestellt. Es sieht so aus, als ob einige Knollen einer ungef?hr S-f?rmigen Kurve folgten, einige wuchsen linear und einige zeigten Perioden des Wachstums, die durch Perioden unterbrochen waren, in denen die Knollen wenn überhaupt nur langsam wuchsen.

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Résumé Des types de tubérisation et de croissance ont été étudiés au cours d'une expérimentation de variétés portant sur Désirée, Maris Piper et Pentland Crown. Plantées à la même densité, ces variétés donnent différentes répartitions de calibre. Pentland Crown tend à produire de gros tubercules; Maris Piper produit un grand nombre de petits tubercules et Désirée est intermédiaire. La tubérisation a été suivie en prélevant des échantillons en cours de végétation 3 fois par semaine et en enregistrant le nombre de stolons et de tubercules formés à chaque noeud (tableau 1). Le grossissement de chaque tubercule, en caisses de bois (fig. 1), et au champ sous polyéthylène (fig. 2), a été suivi en mesurant les trois axes principaux et en transformant ces mesures en volume (on suppose que le tubercule est un ellipso?de), ou en poids (en utilisant la relation linéaire entre produit des axes et poids frais et sec des tubercules cultivés dans des conditions identiques et récoltés chaque semaine). Cette technique donne satisfaction et les valeurs des paramètres pour les premières et dernières mesures de chaque année sont indiquées dans le tableau 2 avec les coefficients de détermination appropriés. Les variétés forment un nombre de stolons similaire mais un nombre de tubercules différents. Des tubercules plus nombreux se sont formés au premier noeud, mais Pentland Crown présentait moins de tubercules aux noeuds, 2–8 que les autres variétés. Le rapport du nombre de tubercules par noeud et du nombre de tubercules formés à chaque noeud, exprimé en pourcentage du nombre de stolons (fig. 3) différait considérablement d'une variété à l'autre. La répartition des calibres de ces variétés a probablement été influencé. L'époque de tubérisation était la même pour chacune d'entre elles. Au cours de cette expérimentation, plus de 500 tubercules ont été mesurés (fig. 4–6). Pour certains tubercules, il appara?t que la courbe tubérisation s'apparente à une sigmo?de. Pour d'autres, la croissance est linéaire, parfois interrompue par des périodes de croissance ralentie.

     

Periderm and lenticel characterization in relation to potato cultivar, soil moisture and tuber maturity

Summary Using the fluorescent lipid stain fluorol yellow 088 to detect suberin in tubers, the numbers of layers and the thickness of the suberized cells in the periderm and within the lenticels showed changes after the early stage of tuber formation. These changes as tubers matured were affected by cultivar and soil moisture conditions. Penetration of the surface tissues of tubers by the water soluble stain safranin O was confined to the lenticles. Although the zone of suberized cells in lenticels acted to some extent as a barrier to penetration, there was not a clear relationship between permeability and suberization. Stored tubers showed thicker suberin berriers in lenticels than freshly harvested tubers. When lenticels of stored tubers proliferated the suberin barrier was disrupted but a further thick suberin barrier formed after exposure to air for two weeks. The findings are discussed in relation to biological control ofErwinia by antagonistic bacteria.     

The effect of stem population on tuber yield in a trial with single-stem seed pieces

Summary Excised eyes on 5 g of seed tuber tissue were used to grow plants with single stems. These were transplanted in the field at densities of 120 000, 180 000 and 240 000 plants/ha, with in-row spacings of 16 or 33 cm. Total tuber yield was not affected, although more tubers were produced at the high densities. Tuber numbers did not increase in proportion to the number of stems. This was due to the fact that the number of tubers per stem decreased from a mean of 4.8 to 2.9 with increasing stem populations.     

The distributions of tuber sizes in droughted and irrigated crops of potato. I. Observations on the effect of water stress on graded yields from differing cultivars

Summary The distributions of the yield and of the numbers of tubers were examined in graded samples of tubers from field experiments designed to compare droughted and irrigated crops of cultivars of potato thought to differ in their tolerance of drought. Drought caused a downward shift of about 7–10 mm in the centres of both distributions. The consequences were that smaller fractions of lower total yields reached the sizes usually specified for table (40–80 mm) and crisping (<45 mm) potatoes. Differences in the distributions of numbers of tubers in droughted and irrigated crops were statistically significant in four of the six cultivars examined. The main reason for lower ware yields in the droughted crops was that fewer tubers reached the minimum size (40 mm). The effect of delayed harvest was to shift the distributions upwards.     

Seed tuber production from true potato seed (TPS) in Egypt and the influence of environmental conditions in different growing periods

Summary The effect of environmental conditions on seed tuber production from true potato seed (TPS) in nurseries was evaluated in different growing periods during two seasons in Egypt. In the autumn season, high soil and air temperatures reduced emergence and initial foliage development of early-sown seedlings, and decreased tuber yields. In late-sown seedlings low night temperatures and short daylengths caused early tuberization, poor foliage development and low tuber yields. In the spring season, low temperatures and short daylengths resulted in a slow initial foliage development and early haulm-tuber competition for assimilates in early-sown seedlings. However, tuber yields were higher than for late-sown seedlings, presumably because net assimilation and assimilate partitioning to the tubers were impaired by above optimum air and soil temperatures in May and June. Slow development of true seed-sown plants caused a high sensitivity towards adverse environmental conditions and would limit yielding ability in Egypt.     

The effect of potato seed tuber age and soil temperature on the phenomenon of non-emergence

Summary In three experiments seed tubers of cvs Jaerla, Bintje and Alpha, stored at 12 °C in the dark, were planted in glasshouses on various dates at soil temperatures varying from 8 °C to 28 °C to study the phenomenon of ‘non-emergence’ with ageing seed tubers. In general the tubers were desprouted before planting. At all plantings cv. Alpha produced normal plants. The cvs Jaerla and Bintje, however, showed high proportions of the ‘little potato’ phenomenon when planted in cold soil, and high proportions of decayed tubers more frequently in warm soil. When using physiologically older seed at later planting dates, both the range of soil temperatures became wider and the proportions of seed tubers experiencing both ‘little potato’ and decay increased.     

Effect of pre-planting temperature and light treatments of seed tubers on potato yield and tuber size distribution

Summary From 1985 to 1987 field experiments were caried out to test the influence of different pre-planting storage treatments on yield and size distribution of potato crops. The storage treatments included 4, 8 and 12°C in the light and in the dark, for different periods before planting. Subjecting the seed tubers to light at low temperatures induced crops to produce a larger proportion of smaller sized tubers while storing seed tubers at higher temperatures in the dark led to an increase of tubers >55 mm. These observations were confirmed in five field experiments with two cultivars at two sites and in three growing seasons.     

The effect of some environmental factors on potato tuber numbers

Summary Five experiments studied the effects of shading, temperature and stolon growth medium on stolon and tuber formation. The lowest numbers of stolons and tubers were produced at the higher and lower nodes. Numbers of primary stolons and tubers were relatively stable across nodes and treatments and variation in total numbers was largely due to changes in numbers of lateral and branch stolons. In the field, 70% shading reduced numbers of stolons and tubers by reducing lateral and branch stolons, while in hydroponics, 45% shading and temperatures at initiation had no effect on tuber numbers. In both environments later planting tended to reduce numbers of stolons and tubers. There were large effects of the physical environment, with compost producing three times as many tubers as dry vermiculite. The potential number of tubers may be determined by the number of stolons and the environmental conditions affecting stolon formation and development.     

Possible mechanisms of size hierarchy among tubers on one stem of a potato (Solanum tuberosum L.) plant

Summary The tubers on one potato stem vary greatly in size and their size distribution may be described by their number and their average size and its variation. These components are mutually and closely related. The variation is partly caused by stolon characteristics, including their date of initiation, position and size. Tuber size is therefore partly determined before tuberization. Tuber-specific factors, such as the position and activity of the tuber initial also play a role, but the date of initiation of an individual tuber is not crucial. Later on, during tuber bulking, the activity of enzymes involved in starch metabolism, the levels of hormones, the mineral composition and the turgor potential are associated with the tuber growth rate. The growth characteristics of individual tubers may also vary since they are exposed to different conditions because of differences in position or growing period.     

Glycoalkaloid accumulation during tuber development of early potato cultivars

Summary Plants of early potato cultivars were sampled at successive harvests during tuber development and individual tubers were analysed for α-solanine and α-chaconine using high-performance liquid chromatography. Mean tuber total glycoalkaloid concentration (α-solanine plus α-chaconine) per plant decreased with time and statistically significant differences between cultivars were observed in the patterns of decline during tuber growth. The mean tuber ratio of α-chaconine to α-solanine increased during tuber development and was also affected by genotype. Total glycoalkaloid concentration for individual tubers of marketable sizes (up to 50 g fresh weight) exceeded the safety limit of 20 mg per 100 g fresh weight for cultivars Home Guard and British Queen but not for Rocket. These differences were attributable to differences in both rates and patterns of glycoalkaloid accumulation during tuber development, although in all cultivars results were consistent with glycoalkaloid accumulation occurring in the expanding peripheral cell layers of tubers for a considerable period after tuber initiation.     

Assessment of heat tolerance in potato (Solanum tuberosum ssp.tuberosum) under controlled environmental conditions: the ‘cutting technique’

Summary The ‘cutting technique’ or ‘tuberisation stimulus’ (TS) technique was investigated as a method of identifying heat tolerant clones ofSolanum tuberosum ssp.tuberosum. TS levels were assessed, using leaf cuttings, for 100 experimental clones growing under heat stress in a controlled environment (35/15°C day/night temperatures, 16 h photoperiod). No correlation was found with the tuber yields of these clones when grown in an irrigated trial in Israel. The relationship between TS levels and tuber production was investigated using early and later maturing cultivars grown with and without heat stress. These experiments showed that the cv. Blanka had high TS levels in all environments and high tuber fresh weights when grown under heat stress. However, the correlation between TS levels and tuber fresh weight for all cultivars was weak (r=0.45). The results indicated that this TS technique is unlikely to be an accurate method of screeningtuberosum clones for heat tolerance.     

The distributions of tuber sizes in droughted and irrigated crops of potato. II. Relation between size and weight of tubers and the variability of tuber-size distributions

Summary Tubers from six cultivars of potato grown in irrigated and droughted conditions were examined to show whether observed differences in grade distributions were simple consequences of differences in yield or were the results of changes in tuber shape or in the relative variability (Rv=σ/μ) of the distributions. Differences in tuber shape, indicated by the relations between tuber size (mm) and weight, were statistically significant but were insufficient to cause measurable changes in the grade distributions. There was a range of values of Rv but drought had no statistically significant effect on Rv and may only affect it by influencing the numbers of tubers set. The main effect of drought on the distribution of tuber sizes was through its effect on total yield and hence on mean tuber size. Yield and number of tubers define a unique tuber-size distribution regardless of the soil moisture stress experienced by the crop.     

Yield, water use efficiency and nitrogen uptake in potato: influence of drought stress

Summary A lysimeter experiment was performed to study the optimal allocation of limited water supply in potatoes. Irrigation regimes equal to 40, 60 and 80% of maximum evapotranspiration (ET) were evenly applied over the crop cycle. Other treatments involved withholding 80 mm of irrigation, based on ET, beginning at each of three designated growth stages (tuber initiation, early and late tuber growth). An irrigated control treatment, restoring the entire ET, was included for comparison. Continuous drought stress reduced photosynthesis as irrigation volumes were reduced. Plant biomass and tuber yield decreased almost proportionally to water consumption, so that WUE was roughly constant. N uptake was highest in the control and in 80% ET treatment. Withholding water during tuberisation severely hindered plant physiological processes and penalized tuber yield. Reductions in photosynthesis, total biomass and yield were the greatest when drought was imposed during tuber initiation. The earliest stress resulted in the lowest WUE and N uptake. A new crop water stress index (SI) was proposed, which combines atmospheric demand for water and canopy temperature.     

The distribution of sclerotia ofRhizoctonia solani Kühn on the surface of the potato tuber

Summary Seed tubers of cv. Majestic harvested from untreated plants in August had fewer sclerotia ofR. solani at the rose than at the heel end. As plant growth ceased in September and tubers matured, sclerotia formed on the rose end so that all regions of the tuber had similar concentrations. In a preliminary experiment in the previous year, presumably influenced by seasonal factors, the onset of maturity was such that the rose end had the highest and the heel end significantly the lowest concentration of sclerotia. In the second season destroying haulms with acid decreased both the time necessary for tuber maturation and the accompanying change of distribution of sclerotia when spraying was done in August but not if it was delayed until September. The results can be explained, in part, by postulating tuber exudates as a source of nutrition for the fungus.

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Zusammenfassung Die Untersuchung verschiedener Lagerposten von Pflanzkartoffelknollen liess vermuten, dass die Verteilung von Sklerotien vonRhizoctonia solani Kühn auf den Knollen nicht zuf?llig ist. Diese Annahme wurde geprüft, indem Knollen aus zwei Feldversuchen untersucht wurden. Im ersten Jahr waren die Pflanzknollen auf natürliche Weise mit Sklerotien infiziert. Im zweiten Jahr wurde jede Knolle vor dem Auspflanzen mit einem Standardinokulum behandelt. Im ersten Jahr wurde das Kraut Mitte September mit S?ure (ca. 11.5% H₂SO₄; 11251/ha) behandelt oder blieben unbehandelt. Der Bestand wurde vier Wochen sp?ter geerntet. Im zweiten Jahr wurde das Kraut an einem von fünf Daten gespritzt oder nicht gespritzt und nachher in Abst?nden geerntet (Tabelle 1). Jede Knolle war in vier gleiche Abschnitte eingeteilt, der erste beim Kronen- und der letzte beim Nabelende, und wurde, wie in Tabelle 2 aufgeführt, auf Sklerotien bonitiert. Im zweiten Jahr wurde durch jede Knolle ein L?ngsschnitt gemacht und der Prozentsatz des Periderms notiert, das — beginnend am Kronenende — mit einer Pinzette entfernt werden konnte. Das Ergebnis wurde als Beurteilung der ‘Reife’ verwendet. Die Ergebnisse des ersten Jahres (Tabelle 3) zeigten, dass ein signifikanter Zusammenhang zwischen dem Knollenabschnitt und dem Grad der Infektion besteht. Das Kronenende wies das gr?sste Infektionsvorkommen auf, das Nabelende signifikant weniger. Die Histogramme in den Abb. 1 und 2 sind für das zweite Jahr und zeigen die Konzentration der Sklerotien auf s?ure-bzw, unbehandelten Parzellen. Die Leichtigkeit, mit welcher das Periderm von diesen Knollen entfernt werden konnte, wird in Abb. 3 dargestellt. Im August geerntete Saatknollen von unbehandelten Pflanzen wiesen am Kronenende weniger Sklerotien auf als am Nabelende. Als das Pflanzenwachstum im September aufh?rte und die Knollen reiften, bildeten sich am Kronenende Sklerotien. so dass alle Knollenteile gleiche Konzentrationen aufwiesen. Durch die Kraut vernichtung mit S?ure nahmen sowohl die für das Reifen der Knollen notwendige Zeit als auch der damit einhergehende Wechsel der Sklerotienverteilung ab, sofern das Spritzen im August geschah, aber nicht, wenn es bis September verschoben wurde. Unterschiede in der Sklerotien verteilung zwischen den Ergebnissen des ersten Jahres und einem entsprechenden Datum der Krautvernichtung und der Ernte im zweiten Jahr dürften saisonbedingten Unterschieden, welche die Reifeverh?ltnisse beeinflussten, zuzuschreiben sein. Die Ergebnisse k?nnen teilweise erkl?rt werden, wenn man voraussetzt, dass Knollenausscheidungen dem Pilz als Nahrungsquelle dienen, die auf einer jungen Knolle das Pilzwachstum anregt, w?hrend deren Entzug bei der Reifung der Knolle Sklerotienbildung verursacht.

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Résumé Un examen de plusieurs lots de tubercules de pomme de terre de semence suggérait que la distribution àleur surface des sclérotes deRhizoctonia solani Kühn n’était pas au hasard. Cettesuggestion a été contrólée en examinant des tubercules récoltés dans deux champs expérimentaux. La première année, les tubercules de semence étaient infectés naturellement par des sclérotes, mais la seconde année un inoculum standard était appliqué à chaque tubercule préalablement à la plantation. La première année, les fanes étaient traitées ou non à l’acide sulfurique (11,5%H₂SO₄: 1125l/ha) à la mi-septembre et la récolte faite 4 semaines plus tard: la seconde année, les fanes étaient traitées ou non à l’une des 5 dates mentionnées dans le tableau 1 et récoltées à intervalles réguliers par la suite. Chaque tubercule était divisé en 4 régions égales, la première étant à la couronne et la dernière au talon et la présence de sclérotes notée conformément au tableau 2. La seconde année seulement. chaque tubercule était coupé selon l’axe longitudinal et on notait le pourcentage de périderme, se trouvant à l’extrémité de la couronne, qui pouvait être enlevé à l’aide de petites pinces; le résultat était considéré comme une méthode d’appréciation de la maturité. Les résultats de la première année (tableau 3) ont montré qu’ily avait une relation significative entre la région du tubercule et le niveau d’infection à sa surface. L’extrémité de la couronne avait la plus grande contamination. l’extrémité du talon significativement moins. Les histogrammes des figures 1 et 2 concernent la seconde année et indiquent la concentration de sclérotes respectivement sur les parcelles traitées et non traitées à l’acide. La facilité avec laquelle le périderme peut être enlevé de ces tubercules est montrée à la figure 3. Les tubercules de semence. récoltés à partir de plantes non traitées au mois d’ao?t, avaient moins de sclérotes à la couronne qu’au talon. Dès que la pousse des plantes cessa en septembre et que les tubercules m?rirent, les sclérotes se formèrent tellement à la couronne que toutes les parties du tubercule eurent des concentrations identiques. En détruisant les fanes avec de l’acide, on obtint à la fois une réduction du temps nécessaire à la maturation des tubercules et une modification de distribution des sclérotes avec la pulvérisation en ao?t mais en la repoussant jusqu’à septembre. Les diffèrences dans la distribution des sclérotes entre les résultats de la première année et une date correspondante de destruction des fanes et de récolte dans la deuxième année peuvent en être attribuées aux différences de saisons, affectant la vitesse de maturation. Les résultats peuvent être expliqués en partie en postulant quel’exudation des tubercules est une source de nourriture pour les champignons, ce qui, sur un jeune tubercule, stimule la croissance des champignons tandis que son absence, lorsqu’un tubercule m?rit, provoque la formation des sclérotes.

     

Variation in the duration of tuber dormancy within a seed potato lot

Summary The variation in duration of dormancy within a seed tuber lot was studied over three years by harvesting individual plants of cvs Diamant and Désirée from field plots and by storing the tubers at 18°C. The variation in dormancy within a tuber lot was large (especially for cv. Diamant) and was mainly caused by variation within plants. For cv. Diamant there was a close negative relation between dormancy and the cube root of tuber weight, whereas for cv. Désirée a relation with tuber weight was almost absent. The duration of dormancy of a seed lot comprising tubers with a narrow range in weight can be well described by two parameters. It is proposed to maintain the moment of 80% sprouting as the criterion for the end of dormancy of a tuber lot and to characterize the spread in dormancy duration by the time lapse between 10% and 90% sprouting.