Studies in the Physiological Action of 2, 2-Dichloropropionic Acid: I.: MECHANISMS CONTROLLING THE INHIBITION OF ROOT ELONGATION

The inter-relationships between time and concentration and thedegree of inhibition of root elongation have been examined forSorghum vulgare, Zea mays, Helianthus annuus, and Pisum sativum.For all species the inhibitory effect is cumulative but thereis a tenfold difference in the concentration required to halvethe elongation of the most sensitive (S. vulgare) and most resistantspecies (P. sativum). From a comparison of the growth of intactsroots and isolated segments, together with estimates of cellnumber, it has been established that the primary effect is tointerfere with meristematic activity in the root tip, wherethe mitotic cycle is arrested at prophase. Using 2, 2-dichloropropionic acid, containing chlorine-36, thecourses of uptake by both intact roots and isolated segmentshave been followes. In every instance uptake is cumulative withthe greatest accumulation in the root tip. There are again largespecific differences but of a reverse order; uptake is leastfor P. sativum and greatest for S. vulgare. For these two speciesand Z. mays, it is concluded that the magnitude of the equi-effectiveconcentration required to halve root elognation is dependenton the level of accumulation rather than on the reaction atcell level: the cells of H. annuus are more sensitive.     

Effect of Verticillium dahliae on Photosynthesis, Leaf Expansion and Senescence of Field-grown Sunflower

On the basis of known sunflower (Helianthus annuus L.) responsesto soil water deficit, it is proposed that the effect of thefungus Verticillium dahliae Klebahn on plant leaf area precedesand is greater than its effect on leaf photosynthesis and stomatalconductance. To test this hypothesis, we measured shoot andleaf area growth, leaf photosynthetic rate, stomatal conductanceand disease symptoms in a field experiment including hybridsof high (Sankol) and low (Dekasol 3900) susceptibility to V.dahliae. Plants inoculated with V. dahliae and controls werecompared. We also investigated the effect of V. dahliae on keycomponents of plant leaf area, leaf expansion and senescence,in inoculated and control plants of Sankol and Toba, a hybridof intermediate susceptibility to V. dahliae. Reduction in plantleaf area caused by V. dahliae was first detected 31 d afterinoculation (DAI), when visual symptoms of disease in inoculatedplants were slight (Sankol) or absent (Dekasol 3900). Reductionin leaf photosynthesis was first observed 66 DAI; stomatal conductanceand leaf dark respiration were both unaffected by V. dahliaeduring the whole experiment. In comparison with controls, V.dahliae reduced seasonal duration of plant leaf area by 25%in Dekalb 3900 and by 55% in Sankol, whereas the average reductionin leaf photosynthetic rate was 9%. In correspondence with thereduction in leaf area duration, inoculation reduced shoot drymatter of mature Sankol by 50%. In both experiments, less leafexpansion accounted for most of the early reduction in plantleaf area; as the disease progressed, increasing senescencealso contributed to reduced plant leaf area. It is concludedthat the response of sunflower to V. dahliae resembled the responseof the plant to soil water deficit: (1) plant leaf area, ratherthan leaf photosynthetic rate, accounted for the reduction ingrowth in mass; and (2) reduced leaf expansion early in theseason and faster leaf senescence in older plants accountedfor the decrease in plant leaf area. Copyright 2000 Annals ofBotany Company Helianthus annuus, Verticillium dahliae, allometry, apical dominance, drought, leaf expansion, leaf senescence, photosynthesis, stomatal conductance, growth     

Physiological and Ecological Studies in the Analysis of Plant Environment: VII. An Analysis of the Differential Effects of Light Intensity on the Net Assimilation Rate, Leaf-Area Ratio, and Relative Growth Rate of Different Species

Since relative growth rate is the product of net assimilationrate and leaf-area ratio (leaf area/plant weight), it followsthat if the effects of shading on both net assimilation rateand leaf-area ratio can be expressed mathematically, then therelationship between light intensity and relative growth ratecan be derived from the product of the two mathematical expressions. For all the ten species investigated in field and pot cultureexperiments, it has been found that during the early vegetativephase both the changes in leaf-area ratio and net assimilationrate, over the range of 0·1 to full daylight, are linearlyrelated to the logarithm of the light intensity. In consequence,the relationship between relative growth rate and the logarithmof light intensity—being the product of the two linearregressions—is curvilinear. For species of shady habitats (Geum urbanum, Solamun dulcamara)neither the levels of assimilation rate nor the ‘compensation-point’values are very different from those of the eight species fromopen situations (e.g. Hordeum vulgare, Pisum sativum, Fagopyrumesculentum). Nevertheless the intensity at which growth rateis maximal varies between species: it is 0•5 for G. urbanum,0•7 for H. annuus, full daylight for F. esculentum, whilefor Trifolium subterraneum the calculated value is 1·8daylight. Such specific differences can be largely accountedfor in terms of the differences in leaf-area ratio at the differentlight levels. On the basis of this analysis of the light factor, a ‘shade’plant is best redefined as a species in which a reduction ofthe light intensity causes a rapid rise in the leaf-area ratiofrom an initial low value in full daylight: for a ‘sun’plant the converse definition holds.     

The Vegetative Growth of Helianthus annuus and Phaseolus vulgaris as Affected by Seasonal Factors in Freetown, Sierra Leone

Sand-culture experiments were carried out in full daylight atsuccessive weekly intervals between March and December 1969,to investigate the effects of seasonal changes in climatic factorson the growth of Helianthus annuus and Phaseolus vulgaris inFreetown. Values for a number of growth parameters were calculatedfrom the dry weights of the leaves, stems, and roots, and fromthe leaf areas. Simultaneously the diurnal changes in climaticfactors were recorded. Multiple regressions linking light, temperature, and relativehumidity with some of the growth parameters were calculated.The total variance accounted for in the regressions of relativegrowth-rate, net assimilation rate, and leaf weight ratio onlight, temperature, and relative humidity ranged from 51 to60 per cent in P. vulgaris. In H. annuus relative humidity wasless important; the percentage proportion of total varianceaccounted for in the regression of leaf weight ratio (and leafarea ratio in both species) on light and temperature was notsignificant. The results showed that H. annuus grew faster than P. vulgaris,but the latter's growth was depressed less by the dull weatherof the rainy season. The relative growth-rates of both specieswere positively dependent on light and temperature while theirnet assimilation rates were negatively dependent on relativehumidity, and their leaf area ratios negatively dependent onlight. All parameters used except leaf area ratio and leaf areato leaf weight ratio showed seasonal variations correspondingto dry and rainy seasons. The initial and final values of leafarea ratio and weight ratios were always different but did notvary in the same direction in both species. The results are discussed in relation to similar work done elsewhere.     

Roles of the Af and Tl Genes in Pea Leaf Morphogenesis: Shoot Ontogeny and Leaf Development in the Heterozygotes

The wildtype leaf blade of Pisum sativum possesses proximalleaflets and distal tendrils, which may be altered by two recessivemutations that affect pinna morphology, afila (afaf) and tendrilless(tltl). Using morphological observations and SEM, the variationin leaf forms along the plant axis and leaf development werecharacterized for plants heterozygous at the Af and/or Tl loci.The Af and Tl genes interacted to affect many characteristicsof shoot ontogeny, including rate changes in leaf blade lengthand complexity increases, as well as time to flowering. TheAf gene retarded early vegetative development and acceleratedthe time to flowering. The leaf phenotypes of these heterozygousgenotypes were specified mainly by changes in the timing ofmajor developmental events. The data support the hypothesesthat both genes are heterochronic in nature and that the pealeaf blade consists of three genetically- and developmentally-determined regions: proximal, distal and terminal. Copyright2000 Annals of Botany Company Heterochrony, leaf development, shoot ontogeny, Pisum sativum L., garden pea, afila,tendrilless .     

Response of Cultured Embryos of Phaseolus vulgaris and Hordeum vulgare to Farnesol

Excised embryos of Phaseolus vulgaris incubated in a mediumcontaining 10 mg dm^–3 farnesol showed enhanced root growthwhereas the leaves remained rudimentary At lower concentrationsof exogenous farnesol normal leaf development occurred and rootgrowth was comparable to untreated cultures. Enhanced root growthalso occurred when excised embryos of Hordeum vulgare were treatedwith farnesol but only at 10 mg dm^–3 and this treatmentdid not prevent leaf growth X-ray micro-probe analysis of leavesrevealed an increased phosphorus content in P vulgaris and adecreased sulphur content in H vulgare in comparison to untreatedplants. Hordeum vulgare L., barley, Phaseolus vulgaris, bean, embryo culture, farnesol, X-ray microprobe analysis, root growth     

Leaf Primordium Initiation and Expanded Leaf Production are Co-ordinated through Similar Response to Air Temperature in Pea (Pisum sativumL.)

Accurate prediction of the timing of leaf area development isessential to analyse and predict the responses of crops to theenvironment. In this paper, we analyse the two processes determiningthe chronology of leaf development—initiation of leafprimordia by the shoot meristem and production of expanded leavesout of the shoot tip—in several pea (Pisum sativumL.)cultivars in response to air temperature and plant growth rate.Contrasting levels of air temperature and plant growth rateduring leaf development were induced by a wide range of sowingdates and plant densities in glasshouse or field experiments.Full leaf expansion was found to occur one phyllochron afterfull leaf unfolding, whatever the leaf nodal position. Primordiuminitiation and expanded leaf production rates presented similarquantitative responses to air temperature (linear response andcommonx-intercept), whatever the plant growth rate, cultivaror period of cycle. As a consequence, they were co-ordinatedand the numbers of initiated primordia or expanded leaves wereeasily deduced from simple visual observation of leaf unfolding.The change, over time, of the numbers of initiated leaf primordiaand fully expanded leaves correlated with cumulated degree-days,with stable relationships in a wide range of environmental conditions.Two phases, with different production rates, had to be considered.These results allowed us to predict accurately the beginningand the end of individual leaf development from daily mean airtemperatures. The relationships obtained here provide an effectiveway of analysing and predicting leaf development responses tothe environment. Pisum sativumL.; pea; number of leaf primordia; number of leaves; temperature; modelling     

Salt Tolerance in the Triticeae: K/Na Discrimination in Barley

Concentrations of ions were measured in the youngest fully-expandedleaves of Triticum aestivum, T. durum, Hordeum vulgare, H. spontaneum,Secale cereale, and Aegilops squarrosa accessions grown in hydroponicculture in the presence of salt (NaCl+CaCl₂). Triticum aestivum,Secale cereale, and Ae. squarrosa had the low leaf Na and highleaf K concentrations typical of plants which contain the enhancedK/Na discrimination character originally found in Ae. squarrosa.T. durum and the Hordeum species did not have this character.The better growth of H. vulgare than of T. durum with similarsalt concentrations in the youngest fully-expanded leaves maybe a result of better compartmentation of Na, Cl, and K betweendifferent tisssues or between different compartments withincells. The enhanced K/Na discrimination character was expressedin disomic addition lines of H. vulgare chromosomes in Triticumaestivum. The H. vulgare variety Herta and its slender mutantboth had similar leaf cation concentrations, although they differedin growth rate when grown at 60 mol m^–3 NaCl. H. vulgareand T. durum seedlings grown in the absence of monovalent cationsaccumulated more ²²Na in their shoots than seedlings of otherspecies when incubated in 1.0 mol m^–3 NaCl labelled with²²Na. Key words: Salt, ion transport, I genome, barley, wheat     

Barbiturate-induced Disorientation of Root and Shoot Geotropism in Germinating Cruciferous Seedlings

Cruciferous seedlings germinated on glass-distilled water andon barbital manifested normal root and shoot geotropisms whereastwisted roots and shoots were evident in seedlings germinatedon amobarbital and secobarbital, the roots often being completelysuspended in the air. The marked difference in germination anddevelopment behaviour of cress and white mustard seedlings germinatedon barbital compared with that on amobarbital and secobarbitalindicate that the Meyer-Overton principle of drug potency andlipid solubility was operative in both species of germinatingseedlings. Lepidium sativum L., Sinapsis alba L., seed germination, geotropism, barbiturates     

Effects of Age and Gibberellic Acid on the Flavonol Content of Pisum sativum Leaves

Changes in the content of quercetin and kaempferol during leafdevelopment in Pisum sativum cv. Meteor are described. As theleaves matured the quercetin content decreased and that of kaempferolincreased during a period from 37 to 48 days from planting.Maximum flavonol content per leaf was found in leaves higheron the shoot as development proceeded, but total leaf flavonolcontent per plant remained relatively constant. There was noevidence to suggest that the quantitative changes in the individualflavonols were the result of conversion of quercetin into kaempferol. Treatment with gibberellic acid led to an accelerated fall inquercetin content in the mature leaves. This change was detectable24 h after treatment and before any apparent growth response.     

Rewatering plants after a long water-deficit treatment reveals that leaf epidermal cells retain their ability to expand after the leaf has apparently reached its final size

Background and Aims: Leaves expand during a given period of time until they reachtheir final size and form, which is called determinate growth.Duration of leaf expansion is stable when expressed in thermal-timeand in the absence of stress, and consequently it is often proposedthat it is controlled by a robust programme at the plant scale.The usual hypothesis is that growth cessation occurs when cellexpansion becomes limited by an irreversible tightening of cellwall, and that leaf size is fixed once cell expansion ceases.The objective of this paper was to test whether leaf expansioncould be restored by rewatering plants after a long soil water-deficitperiod. Methods: Four experiments were performed on two different species (Arabidopsisthaliana and Helianthus annuus) in which the area of leavesthat had apparently reached their final size was measured uponreversal of water stresses of different intensities and durations. Key Results: Re-growth of leaves that had apparently reached their finalsize occurred in both species, and its magnitude depended onlyon the time elapsed from growth cessation to rewatering. Leafarea increased up to 186% in A. thaliana and up to 88% in H.annuus after rewatering, with respect to the leaves of plantsthat remained under water deficit. Re-growth was accounted forby cell expansion. Increase in leaf area represented actualgrowth and not only a reversible change due to increased turgor. Conclusions: After the leaf has ceased to grow, leaf cells retain their abilityto expand for several days before leaf size becomes fixed. Aresponse window was identified in both species, during whichthe extent of leaf area recovery decreased with time after the‘initial’ leaf growth cessation. These results suggestthat re-growth after rewatering of leaves having apparentlyattained their final size could be a generalized phenomenon,at least in dicotyledonous plants.     

Changes in Poparity of Growth During Leaf Initiation in the Pea, Pisum sativum L.

In the apical dome of the pea shoot apex the axis of growthof the epidermal cells becomes predominantly longitudinal inthe I₁ region one plastochron before a leaf is initiated, andthis orientation persists into the young primordium. In contrast,in the underlying, non-epidermal cells the growth axis in theI₁ region becomes randomized half a plastochron before leafinitiation, but in the young primordium it becomes the sameas in the epidermis. The initiation of a leaf primordium thereforetakes place without any major change in the orientation of growthaxes in the epidermis. A controlling role for the epidermisis therefore suggested. No marked reorientation of the growthaxis occurs on the sides of the newly initiated primordium.The shape of the young primordium can be related to the differentialrates of growth and division within it rather than to changesin growth orientation. Pisum sativum, pea, shoot apex, meristem, growth, epidermis, polarity     

Does ABA in the Xylem Control the Rate of Leaf Growth in Soil-Dried Maize and Sunflower Plants?

Maize (Zea mays L.) and sunflower (Helianthus annuus L.) plantswere grown in large volumes of soil and leaf growth rate wasmonitored on a daily basis. Half the plants were given a soildrying treatment and when they showed a significant restrictionof growth rate (compared to both their daily growth rate beforedrying and the average growth rate of well-watered plants onthe same day), leaf water relations were measured and xylemsap was extracted using several techniques. There was a significant negative log-linear relationship betweenthe rate of leaf growth and the concentration of ABA in thexylem for both species. There was no clear relationship betweenleaf growth rate and leaf water potential or turgor for eitherspecies. Assessment of different methods for sampling xylemsap suggests that exudates collected from stem stumps or samplescollected by pressurizing the whole root system are suitablefor estimating ABA concentration in xylem, at least with largeplants of maize or sunflower, provided the first few hundredcubic millimetres of collected sap are used for the assay. Centrifugationof sections of stems resulted in dilution of ABA in the xylemsap with sap squeezed from parenchyma tissue. This is because,at least in plants subjected to mild soil drying, the concentrationof the ABA in the xylem is far higher than that in the cellsap of stem tissue. Results support the proposal that ABA plays a major role asa chemical signal involved in the root-to-shoot communicationof the effects of soil drying. The non-hydraulic restrictionof leaf growth by a chemical signal can be explained by theextra root-sourced ABA in the xylem and may be an importantcomponent of the modification of growth and development whichresults from prolonged soil drought. Key words: Soil drying, ABA, leaf growth, Zea mays L., Helianthus annuus L.     

Genetic Control of Leaf Morphology: A Partial View

The partial-shoot theory of the leaf was a controversial hypothesisrevived by Arber and supported by her morphological and anatomicalstudies. This theory highlighted the parallels between leavesand shoots and contrasted with an alternative view that leaves,with their limited growth potential, are completely distinctfrom shoots. Pea morphological mutants with altered growth potentialin their compound leaves are described. The unifoliata mutanthas a limited growth potential relative to wild-type;cochleata,afila and insecatus have extended potentials. Characterizationof theunifoliata mutation and gene expression patterns indicatethat unifoliata is a common factor in pea compound leaf andfloral shoot development, and so provides rudimentary supportfor the idea that some leaves have shoot-like characteristics.Tomato leaves are also considered to lend tentative support.The afila and insecatus leaf forms are described as bipinnateand weakly bipinnate, respectively. These and the tendril-lessmutant are potential phenocopies of legume relatives, an ideabased on Vavilov's law of homologous series of variation. Arberillustrated, but did not articulate in genetical terms, thatmorphological variation in structures within an individual plantcan be interpreted as reiteration of design. Analogous withVavilov's view, this can be considered a consequence of thesame genetic programme in a different location.Copyright 2001Annals of Botany Company Agnes Arber, compound leaf, genetics, knotted1, legume, morphology, mutant, partial-shoot, pea, Pisum sativum, unifoliata, Vavilov     

Evaluation of a Non-Destructive Method for Measuring Turgor Pressure in Helianthus

The portable instrument described by Heathcote, Etherington,and Woodward (1979) for the non-destructive measurement of turgorpressure was evaluated in Helianthus annuus and Helianthus paradoxus.A good correlation was obtained between turgor pressure measuredwith the instrument and turgor pressure estimated by the pressure-volumetechnique for individual leaves allowed to dry after excision;however, variation in both the intercept and slope of the relationshipoccurred between leaves. Consequently, there was no correlationbetween the output of the instrument for individual leaves andthe turgor pressure of the same leaves estimated by conventionalmethods. Moreover, for a given leaf, the instrument had onlya limited ability to detect temporal variation in turgor pressurewhen compared with turgor pressure calculated from measuredvalues of leaf water potential and leaf osmotic potential. Theinstrument's output was influenced by its proximity to majorveins and by leaf thickness. We conclude that variability inleaf thickness and the presence of large veins limits its usefulnessfor measurement of turgor pressure in Helianthus. Key words: Leaf thickness, Turgormeter, Turgor pressure, Helianthus     

Adaptation and Functional Balance Between Shoot and Root Activity of Sunflower Plants Grown at Different Root Temperatures

Helianthus annuus L. plants were grown with the shoots at normalair temperature and with the roots in nutrient solution at 10,20 or 30 °C. The higher the root temperature the greaterthe growth of the leaves, resulting in higher production ofphotosynthates. Irrespective of growth conditions an equilibriumwas established between the maintenance respiratory activityof shoot and roots. A constant proportion of the photosynthateproduced was used in respiration. The results are discussedin relation to a thermodynamic theory of stability of biologicalsystems. It is suggested that changes in energy partition betweenmaintenance and growth, and then in relative growth rates ofshoots and roots during an adaptation period, represent a majorhomeostatic mechanism. shoots, roots, maintenance respiration, growth, relative growth rate, respiration, adaptation, sunflower, Helianthus annuus L.     

Growth changes in the shoot apex of Sinapis alba during transition to flowering

The aim of the work was to report morphological changes whichoccur in the shoot apex during the morphogenetic switch to floweringin the model long day (LD) plant, Sinapis alba. During the floraltransition induced by 1 LD the growth rate of all componentsof the shoot apex is modified profoundly. The earliest changes,detected at 24 h after start of LD, include a decrease in plastochronduration and an increase of growth of leaf primordia. One daylater, the meristem dome starts to increase in volume, apicalinternodes have an increased height and there is a precociousoutgrowth of axillary meristems. All these changes precede initiationof flower primordia, which starts at about 60 h after the startof LD. Later changes include meristem doming, a decrease inthe plastochron ratio and a shift to a more complex phyllotaxis.All the changes, except the decreased plastochron ratio, arecharacteristics of an apex with an increased tempo of growth.The stimulation of longitudinal growth (height of apical intemodes)is more marked and occurs earlier than the reduction of radialgrowth (plastochron ratio). Key words: Axillary meristem, internode growth, leaf growth, plastochron ratio, plastochron duration     

The Physio1ogical Activity of 2 : 6-Substituted Phenoxyacetic Acids

Anumber of 2 : 6-substituted phenoxyacetic acids in which theside chain had been substituted by ailcyl groups to form -propionic,-n-butyric and -n-valeric acids were investigated to assesstheir ability to act as growth-regulators. Besides employingstandard tedmiques of bioassay, further experiments were undertaken to determine the effects of these compounds on the vegetativegrowth and development of intact plants of Helianthus annuus. It has been established that all the compounds tested inducedcurvature in the Went pea curvature test and that without exceptionthe activity of the parent phenoxyacetic acid (2:6-dichloro-,2:4:6-trichloro-,2:6-dimethyl-, and 2:4-dichloro-6-methyl-)was increased by side chain substitution. In the Avena straight growth assay, the a 2:6-dichloro- and2:6-dimethyl- phenoxypropionic and butyric acids brought aboutstatistically significant increases in length of the coleoptilesections, when measurements were made at the end of 24 hours.Their activity was of the same order as that exhibited by a2:4-dichlorophenoxyacetic acid. Investigations were carried out on the uptake of water by sectionsof pea internode and the extension growth of Avena coleoptileaectiona when both the concentration and the length of treatmentwere varied. For some compounds it was found that over a certainrange of concentration water uptake and extension growth wereaccelerated in the initial 6–8 hours. Subsequent to this,inhibition occurred so that no significant increases above controlvalues were apparent at the end of 24 hours, and in some casesan actual loss of water or shrinkage in length took place. Itwas thus possible to demonstrate that 2:6-dimethylphenoxyaceticacid acts as a growth regulator in pea extension growth andthat 2:6-dichiorophenoxyacetic acid is active in the Avena test. The changes in the leaf area of individual pairs of leaves,together with the lengths of the internodes and shoot weightwere followed after application of measured amounts of eachcompound to the first pair of leaves of H. annuus. A numberof the 2:6-compounds were capable of modifying growth in wayssimilar to 2:4-dichlorophenoxyacetlc acid. However, much higherconcentration had to be applied in order to produce effectscomparable to those caused by the 2:4-dichloro- compound. Measurementsof the percentage of applied compound which penetrated intothe leaf showed that there were no marked differences in thisrespect between the different phenoxy acids. On the other hand,experiments in which the treated leaves were left on the plantfor varying periods of time led to the conclusion that the a: 6-substituted compounds are less readily translocated than2:4-dichlorophenoxyacetic acid.     

Differences in Architecture and Shoot Growth during Stagnant and Extension Growth Phases of Acanthopanax sciadophylloides(Araliaceae)

The shoot growth of a deciduous tree, Acanthopanax sciadophylloidesFranch. et Savat. shows inter-annual intermittent repetitionof two distinctive phases, a stagnant growth phase (S-phase)and vigorous extension-growth (E-phase). To help understandthe differentiation mechanism, shoot development was studiedover time in both shoot phases. S-phase and E-phase shoots weredistinguished from each other by morphological traits: S-phaseshoots are characterized by higher allocation to leaves anda shorter period of stem growth, while E-phase shoots show continuousstem extension over the growing season. Specific leaf area didnot differ between the two phases. This shoot differentiationwas similar to the morphological differentiation of shoots betweenlong vs. short shoots found in some temperate trees. Leavesof both phases were well-dispersed through adjustment of petiolelength and leaf-blade size to reduce mutual shading within ashoot. Stem-wood density of current-year shoots was lower inE-phase compared with S-phase shoots. Leaves produced earlyin the season affected the growth phase of the following year.These results suggest that annual shoot differentiation of A.sciadophylloides was determined during the previous season andreflects leaf productivity in a given habitat during that growingseason. Copyright 2001 Annals of Botany Company Acanthopanax sciadophylloides, Araliaceae, biomass allocation, intermittent shoot growth, leaf display, shoot architecture, shoot differentiation