Flowering in Pisum: the Effect of Light Quality on the Genotype If e Sn Hr

Far-red light, when given as a 16 h photoperiod extension, iamore effective than red light in reducing the flowering nodeof genotype Pisum. In contrast, when a 16 h dark period is interruptedby a 2 h light break red light is more effective than far-redlight. In addition, the stimulatory effect of a red interruptionis partially reversed by a subsequent period of far-red. However,a light interruption is not effective until over 12 h have elapsedsince the start of the previous photoperiod, regardless of whetherthe photoperiod was of 4 or 8 h duration. The results suggest that there are two light-dependent reactionscontrolling flowering in peas, one operating through the phytochromesystem with high levels of Pfr suppressing production of flowerinhibitor by the sn gene and a second requiring continuous illuminationwith wavelengths above 700 nm. The role of time measurementin the photoperiod response in peas is suggested to be filledby the proportion of time the Sn gene is effectively producinginhibitor. The photoperiod response in peas is not independentof temperature or plant age since the activity of gene Sn isalso varied by these factors.     

Flowering in Pisum: the Sites and Possible Mechanisms of the Vernalization Response

Grafting experiments with several genotypes provide evidencethat vernalization acts through at least two mechanisms. Vernalization of the stock promoted flowering by 26 nodes ingenotype If e Sn Hr and 5.5 nodes in genotype If e Sn hr buthad no detectable effect in genotype If e sn hr. Cold treatmentappears to cause a higher ratio of promoter to inhibitor, atleast in part, through low temperature repression of Sn activity.This mechanism is particularly evident in the cotyledons sincethey form a major area of Sn activity during vernalization.Continuous light was shown previously to prevent Sn forminginhibitor. It seems therefore that both photoperiod and vernalizationhave an effect through the Sn gene. Vernalization of the shoot promoted flowering by 19 nodes ingenotype If e Sn Hr, 3 nodes genotype If e Sn hr, and 1 nodein genotype If e sn hr1 grafted to an If e Sn hr stock. Theshoot effect may result from one or possibly two mechanisms.Firstly, vernalization may lower the threshold ratio of promoterto inhibitor required at the apex for floral initiation. Thesame change in threshold could result in changes in the floweringnode of quite different magnitude depending on the rate of changein the hormonal levels in the different genotypes. Secondly,vernalization may disturb the ageing process relative to theplastochronic age leading to an earlier (nodewise) decline ininhibitor level.     

Flowering in Pisum: the Effect of Age on the Gene Sn and the Site of Action of Gene Hr

Late cultivars of peas behave as quantitative long day plants.The reason that they flower between nodes 20 and 35 under an8 h photoperiod is shown to be because the leaves and maturestem produce a more promotory ratio of the flowering hormonesas they age. Later formed leaves may also start with a slightlymore promotory ratio than the leaves produced at a lower node.The gene Sn controls the production of a flower inhibitor andit is suggested that the activity of this gene in a leaf isgradually reduced as the leaf ages. From grafting experiments,the site of action of the gene Hr is shown to be in the leavesor mature stem and not at the shoot apex. This supports a previoussuggestion that the gene Hr is a specific inhibitor of the ageingresponse of gene Sn. Gene Hr is shown to cause a substantial delay in the floweringnode of decotyledonized plants of genotype If e sn hr undershort day conditions, suggesting that Hr has little effect inthe cotyledons. It is argued that the gene sn is a leaky mutantand that gene Hr does not control a photoperiod response inits own right but has its effect through the Sn locus. From a comparison of intact plants and self-grafts of the lategenotype If e Sn hr it is shown that under the conditions usedphysiological age may be of more importance than chronologicalage in determining flowering in peas. Reasons for the smalleffect of defoliation treatments on flowering are discussedas well as possible reasons for the promotory effect of decotyledonizationon the flowering node of late lines. Pisum sativum L, flowering, ageing, genetic control     

Genotype Differences in Photoperiodic Sensitivity and Vernalization Response in Wheat

The regression of final leaf number on leaf number at transferfrom an 8 h to an 18 h photoperiod was used to compare the photoperiodicresponse of eight locally-adapted lines of wheat. Comparisonsof such regressions in the vernalized and univernalized conditionsenabled detection of the presence or absence of a significantcultivar vernalization response and comparisons of differingresponses between cultivars. Prior vernalization generally did not significantly alter the‘rate’ of photoperiodic response, as the slope ofthe LNT/FLN regression, indicating a certain physiological independenceof vernalization and photoperiodic responses in wheat. Differences, both in photoperiodic and vernalization responseof the eight wheats studied have been discussed in terms ofadaptability and breeding for maturity alteration in wheat.Evidence has been produced for the possible existence of a thirdfactor influencing developmental processes associated with floweringin wheat.     

Flowering in Pisum: the Effect of Genotype, Plant Age, Photoperiod, and Number of Inductive Cycles

The genotypes If e Sn hr, Lf e Sn hr, and If e Sn Hr requirefewer inductive cycles as they age. It is suggested that thisresults from a decrease in the activity of the Sn gene in theleaves as they age, resulting in a higher ratio of promoterto inhibitor. Gene Lf does not affect the rate of this agingbut it does increase the number of inductive cycles requiredfor flower induction over the first 5 weeks of growth. The geneHr has no effect until week 4 but thereafter causes a reductionin the effect of age on the Sn gene. The genotype If e Sn Hrcan be induced by a single inductive cycle (32 h of light) fora relatively long period. The length of dark period required for the expression of theSn gene is shown to be less than 4 h providing a relativelylong photoperiod precedes the dark period. It appears that noper manent induction of tissue by photoperiods favourable toflowering occurs in peas. The critical photoperiod for plantsof genotype if e Sn Hr is shown to be between 12 and 14 h atl7·5 °C and the usefulness of the term ‘criticalphotoperiod’ is discussed with respect to quantitativelong-day plants.     

Flowering in Pisum: Effect of the Parental Environment

The parental photoperiod appears to have no effect on the floweringnode of the progeny, provided the progeny seed is selected tobe of the same weight. However, the parental photoperiod doesinfluence the mean seed weight and seed weight is shown to causesignificant alterations in the flowering node and time as wellas causing alterations in vegetative characters like the internodelength and the rate of leaf expansion and node formation. Theeffect of seed weight on flowering is thought to result fromthe alterations in the growth rate. On the other hand, vernalization of the parents does appearto cause a small, but significant effect on the flowering nodeof the progeny (i.e. it could be transmitted through a meioticdivision). This effect disappears in the next generation andthe possible nature of this effect is discussed. Pisum sativum L, garden pea, flowering, photoperiod, vernalization     

Quantitative effects of the genes Lf, Sn, E, and Hr on time to flowering in pea (Pisum sativum L.)

Flowering time in pea (Pisum sativum L.) is determined by genetically controlled responses to photoperiod and temperature. To investigate these responses, 11 lines homozygous for the flowering genes Lf, Sn, E, and Hr were grown under contrasting semi-controlled photothermal environments and the durations (d) from sowing to first flower (f) were recorded. The effects of the four genes were quantified using a two-plane photothermal model which linearly relates the rate of progress from sowing to flowering (1/f) with the mean pre-flowering values of temperature (T) and/or photo-period (P), based on 1/fa + bT (when P is longer than the critical photoperiod, Pc) and 1/fa + bT + cP (when P<Pc). The main effect of Lf alleles was on temperature sensitivity (b) when P>Pc, which increased in the sequence Lf^d<Lf< lf<lf^a. Gene Hr, when together with Sn, increased photoperiod sensitivity (c) and reduced the intercept (a) when P<Pc. Allele sn determined a single plane response to temperature alone (i.e. a day-neutral response). Gene E, when present with lf Sn, increased 1/f in both the thermal (P<Pc) and photothermal (PPc) domains, mainly by increasing a and b, respectively. Variations in the coefficients of the thermal and photothermal responses determined that the critical photoperiod varied with temperature in all photoperiod-sensitive genotypes. A common base temperature of 0.2C was determined amongst Day-Neutral Class genotypes (sn) and thermal time from sowing to flowering increased in the sequence lf^a<lf< <:f<Lf^d. Intra-Class variations attributed to the Lf alleles were also detected in the Late (Sn hr) and Late High Response (Sn Hr) Classes. The linear photothermal model provided a sound basis for studying the quantitative effects of flowering genes in pea.     

Flowering in Pisum: A Fifth Locus, Veg

In addition to the known loci, If, e, sn and hr, a fifth locus,veg, is shown to control flowering in peas. Regardless of thegenotype for the other flowering genes, plants homozygous forthe gene veg did not initiate flower buds under a wide rangeof photoperiod and temperature regimes, including those normallyhighly promotory in peas. Treatment with various plant growthsubstances and grafting to stocks known to promote floweringalso failed to cause initiation. Gene veg prevented expressionof allelic differences at the If locus but segregation for allelesat the sn and hr loci was clearly visible by examination ofseveral vegetative characteristics. For example, sn hr veg andSn hr veg plants showed an opening of the apical bud, productionof lateral branches, and a reduction in growth rate, leafletsize, internode length and stem thickness at approx the sametime as sn hr Veg and Sn hr Veg plants carrying the Lf allelecommenced fruit production, respectively. The graft-transmissibleinhibitor controlled by gene Sn is therefore not specific forthe transition from vegetative to reproductive growth. Geneveg allows the processes leading to apical and foliar senescenceto be examined independently of any effect of flowering andfruiting. We found that gene Sn influenced the total numberof leaves expanded in veg plants but not the time of shoot senescence,which, in plants without flowers and fruits appeared to resultfrom failure of the root system. Pisum sativum L., garden pea, flowering, senescence, genetics     

Flowering in Pisum: A Sixth Locus, Dne

A sixth major flowering gene, dne, is identified in the gardenpea (Pisum salivum L.). Linkage tests show dne is located onchromosome 3 near locus st. The ability to respond to photoperioddepends on the joint presence of the dominant genes Sn and Dnewhich together confer a long day habit. Genotypes Sn dne, snDne and sn dne are all essentially day-neutral although by examiningseveral flowering criteria under strictly controlled conditionssome small responses could be demonstrated. Like sn, dne reducedthe response to vernalization when substituted into genotypeSn Dne. It is suggested genes Sn and Dne both control steps in a biosyntheticpathway which leads to the production of a graft-transmissibleinhibitor of flowering and apical senescence. Stocks of genotypeSn dne and sn Dne promoted flowering in Sn Dne scions whileSn Dne stocks delayed flowering in Sn dne (and sn Dne) scions.However, reciprocal grafting between genotypes Sn dne and snDne gave no evidence of physiological complementarity correspondingto the genetic complementarity ofSn and Dne. Initial resultssuggest dne may be less effective than sn at blocking inhibitorproduction but this requires confirmation.     

Natural Variation of Flowering Time and Vernalization Responsiveness in Brachypodium distachyon

Dedicated bioenergy crops require certain characteristics to be economically viable and environmentally sustainable. Perennial grasses, which can provide large amounts of biomass over multiple years, are one option being investigated to grow on marginal agricultural land. Recently, a grass species (Brachypodium distachyon) has been developed as a model to better understand grass physiology and ecology. Here, we report on the flowering time variability of natural Brachypodium accessions in response to temperature and light cues. Changes in both environmental parameters greatly influence when a given accession will flower, and natural Brachypodium accessions broadly group into winter and spring annuals. Similar to what has been discovered in wheat and barley, we find that a portion of the phenotypic variation is associated with changes in expression of orthologs of VRN genes, and thus, VRN genes are a possible target for modifying flowering time in grass family bioenergy crops.     

Temperature Response of Vernalization in Wheat: A Developmental Analysis

The vernalization response of wheat ( Triticum aestivum L.)was reinterpreted from a developmental perspective, using currentconcepts of the developmental regulation of wheat morphologyand phenology. At temperatures above 0 °C, the effects ofthe process of vernalization per se in wheat are confoundedby the effects of concurrent vegetative development. These effectsare manifested by differences in the number of leaves initiatedby the shoot apex prior to floral initiation, which in turnaffects the subsequent rate of development to ear emergenceand anthesis. Leaf primordia development during vernalizationand total leaf number at flowering were used to develop criteriato define both the progress and the point of saturation of thevernalization response. These criteria were then used to reinterpretthe results of Chujo ( Proceedings of the Crop Science Societyof Japan 35 : 177–186, 1966), and derive the temperatureresponse of vernalization per se for plants grown under saturatinglong day conditions. The rate of vernalization increased linearlywith temperature between 1 and 11 °C, such that the timetaken to saturate the vernalization response decreased from70 d at 1 °C to 40 d at 11 °C. The rate declined againat temperatures above 11 °C, and 18 °C was apparentlyineffective for vernalization. Total leaf number at saturation,however, increased consistently with temperature, as a resultof the balance between the concurrent processes of leaf primordiuminitiation and vernalization. Total leaf number at saturationincreased from 6 at 1 °C to 13.3 at 15 °C, which inturn influenced the time taken to reach ear emergence. The advantagesof using this developmental interpretation of vernalizationas the basis for a mechanistic model of the vernalization responsein wheat are discussed. Triticum aestivum L.; wheat; vernalization; rate; temperature; primordia; leaf number; flowering     

Genetic Regulation of Flowering in Grafts on Pisum sativum L

Lf, E, Sn, and Hr are major loci that condition the flowering and photoperiod responses of Pisum sativum L. Genetic lines containing the dominant alleles of these loci are characterized by flowering in long days, but having a prolonged (>50 node) vegetative phase in short days. A representative of this class, response type G, was used as a receptor in short days for donors of other flowering response types. The qualitative and quantitative flowering response of G receptors depended on the genotype of the donor. Donors containing sn hr induced the earliest development, followed by sn Hr and Sn hr donors. The Lf and E loci in foliar donors apparently did not affect flowering of G. Five-leaved > single-leaved > cotyledonary donors in effecting a flowering response in G, in part due to the longer life of the foliar donors. The responses of G to the various donors were generally consistent with the proposed roles of Lf, E, and Sn, but the role of Hr in these grafts was unclear.     

Apical Senescence in Pisum: A Direct or Indirect Role for the Flowering Genes ?

Apical senescence was examined in a range of intact and defloweredflowering genotypes under both long and short photoperiods.The flower inhibitor produced by the gene Sn, appears to havea direct effect on apical senescence since it can delay apicalsenescence under short day conditions in the absence of flowerand fruit development or where the rate of such developmentis the same in different treatments. Gene Hr can magnify thiseffect. Gene E, on the other hand, appears to influence apicalsenescence only indirectly through the effect it has on flowerand fruit development. The flowering genes at the If, sn andhr loci are also thought to have indirect effects on apicalsenescence. Even in deflowered plants apical senescence appearsto occur eventually in continuous light in all genotypes testedindicating that the presence of developing fruits, althoughpromotory, is not essential for apical senescence. Pisum sativum L., garden pea, flowering, senescence     

Effects of Temperature, Photoperiod and Seed Vernalization on Flowering in Faba Bean Vicia faba

Factorial combinations of three photoperiods (10, 13 and 16h), two day temperatures (18 and 28 °C) and two night temperatures(5 and 13 °C) were imposed on nodulated plants of six diversegenotypes of faba bean (Vicia faba L.). Plants were grown inpots in growth cabinets from both vernalized (1.5±0.5°C for 30 d) and non-vernalized seeds. The times from sowingto the appearance of first open flowers (f) were recorded. Seedvernalization decreased the subsequent time taken to flowerin almost all genotype x growing environment combinations (theexceptions were plants of the cv. Maris Bead grown in threecooler, short-day regimes). The influence of temperature andphotoperiod on the rate of flowering was quantified, using amodel applied previously to other long-day species of grainlegume in which positive linear relations between both temperatureand photoperiod and the rate of progress towards flowering areassumed to apply. A significant positive linear response ofrate of progress towards flowering to limited ranges of meandiurnal temperature was detected in all six genotypes, but inthree genotypes (Syrian Local Large, Aquadulce and Maris Bead)the 28 °C day temperature reduced the rate of progress towardsflowering - suggesting that the optimum temperature for floweringin these genotypes is below 28 °C. In four genotypes (MarisBead, Giza-4, Aquadulce and BPL 1722) a significant positiveresponse to photoperiod, typical of quantitative long-day plants,was observed only in plants grown from vernalized seeds. Incontrast, plants of the genotype Zeidab Local grown from bothnon-vernalized and vernalized seeds showed the same positiveresponse to photoperiod, whereas plants of the land-race SyrianLocal Large were consistently unresponsive to photoperiod. Theimplications of this range of responses amongst diverse genotypesare discussed in relation to screening germplasm. Vicia faba, faba bean, flowering, photoperiod, temperature, seed vernalization, germplasm screening     

Photothermal Time for Flowering in Lentils (Lens culinaris) and the Analysis of Potential Vernalization Responses

Two cultivars of lentils, Laird and Precoz, were subjected to18 potentially vernalizing treatments, comprising constant temperaturesof 1, 5 or 9 °C in factorial combination with photoperiodsof 8 or 16 h for 10, 30 or 60 d. These seeds or seedlings, togetherwith non-vernalized seeds (as controls), were then transferredto four different growing regimes (‘day’/‘night’temperatures of 18/5 °C or 24/13 °C, factorially combinedwith photoperiods of 11 or 16 h). Variation in the number ofdays from sowing to first flower (f) in the growing regimesfor the controls conformed to the equation I/f = a+b+cP, whereis mean temperature (°C), P is photoperiod (h) and a, band c are genotype-specific constants. Accordingly, when theenvironment varies during development, the photothermal timerequired to flower in day-degrees (°C d) is given by 1/babove a base temperature defined as —(a+cP)/b. Most variationin time to flower could be accounted for by the photothermaltime accumulated in the two successive environments. Therefore,there was no evidence of a specific low-temperature vernalizationresponse in either cultivar. Neither was there evidence of ‘short-day’vernalization, i.e. advancement of flowering resulting frompreliminary short-day treatments. A potential error inherentin the predictive model described arises because it ignoresthe presence of a pre-inductive, photoperiod-insensitive phase;but agro-ecological considerations suggest that this error maynot be important in practice. Lens culinaris, lentil, flowering, photoperiodism, vernalization, photothermal time, screening germplasm