Growth Hormone Changes in Chrysanthemum morifolium: Effects of Environmental Factors Controlling Flowering

Simultaneous quantitative analyses have been made of the endogenouslevels of auxin- and gibberellin like substances, growth inhibitors,and auxin-oxidizing enzyme activity in the cold-requiring Chrysanthemummorifolium cv. Sunbeam subjected to different daylength, lightintensity and temperature regimes known to affect flowering.While little hormone or enzyme activity was found in extractsfrom unvernalized plants, a striking rise in auxin-oxidizingenzyme activity occurred rapidly after the end of cold treatment.Increased auxin activity was also recorded shortly after vernalization.At 28 °C both enzyme and auxin activity declined over aperiod of 3–4 weeks; at 20 °C this response was delayed.Gibberellin activity at 28 °C rose steeply about 2 weeksfrom vernalization and declined several weeks later; at 20 °Ca similar response was less marked. Low light intensity treatment,which may have increased endogenous auxin levels, or exogenousauxin application reduced gibberellin-like substance levelsand cause d devernalization.Phosphon D treatment also loweredgibberellin levels and prevented flowering. An extract fromvernalized plants containing gibberellin-like substances intensifiedthe flowering of partially vernalized test plants. Persistenceof high auxin activity in vernalized plants on long days wasassociated with failure to form normal flower buds. Stem elongationrates correlated in general with levels of endogenous auxin-and gibberellin-like substances. Significant amounts of an abscisin-likeinhibitor were found in extracts of flower buds. The mechanismof natural devernalization is discussed in relation to theseobservations.     

Studies in Vernalisation of Cereals: XV. After-effects of Temperature and Drying during Seed Ripening and the Origin of the Vernalisation Requirement in Petkus Winter Rye

Experiments are described which were designed to test whetherthe unvernalised condition of winter rye grain produced by fullyvernalised parent plants is due to de-vernalisation occurringduring maturation and ripening of the seed. The results showclearly that drying of the grain does not account for the un-vernalisedcondition. Parent ears exposed from the time of their emergenceto a range of non-vernalising temperatures (12–20°C), i.e. including ‘neutral’ levels which are neithervernalising nor de-vernalising, still produced unvernalisedgrain. Even plants raised from grain which had been developedat mildly vernalising temperatures (9° C) benefited fromfurther cold treatment applied during their early growth. Itappears, therefore, that the vernalisation requirement in winterrye arises very early in the formation of the seed, i.e. possiblyat meiosis or fertilisation, and that there is no subsequenttransmission of the vernalisation stimulus from the parent plant. Additional observations on plant height, ear size, &c.,also indicate marked effects of temperature and daylength treatments.     

Factors controlling Flowering in the Chrysanthemum: III. FAVOURABLE EFFECTS OF LIMITED PERIODS OF LONG DAY ON INFLORESCENCE INITIATION

Experiments are described which indicate that Chrysanthemumcuttings derived from unvernalized long-day stock plants flowersooner and with lower leaf numbers after vernalization thansimilarly treated cuttings from short-day stock. Long-day treatmentof young cuttings also hastens inflorescence initiation providedthe period of such treatment is limited and given before orimmediately after vernalization. The effect of long day appearsto be maximal when vernalization is complete. Long-day treatmentcannot substitute for vernalization.     

Effects of Photoperiod on the Apparent Viscosity of Leaf Cytoplasm in Kalanchoe blossfeldiana. I. A Method for Quantitative Estimation and its Sources of Error

An adaptation of the centrifuge method of determining cytoplasmicviscosity is described, suitable for the examination of leavesof higher plants exposed to different photoperiodic conditions.Leaf segments are centrifuged for several standard periods,usually i, 2, and 4 minutes, followed by immediate fixation,microscopic examination of hand-microtome sections, and thedetermination of the proportions of cells in the mesophyll regionwhose chloroplasts have been displaced. Among factors whichmay cause systematic errors, variations in cell size, chloroplastsize and chloroplast starch content have been found to causerelatively little error. Similarly, changes in water contentwhich cause up to 14 per cent loss of fresh weight have no largeor consistent effect. Significant effects may arise from diurnalchanges, or from differences in ambient and in growth temperature,as well as in leaf age and the region within the leaf that istested; they can all be minimized by appropriate sampling techniques.Any treatment effects revealed by the method are clearly notascribable to viscosity differences alone, but they are likelyto represent real changes in the physical state of the cytoplasm;they are therefore referred to as representing changes in ‘apparentviscosity’.     

Growth and Dormancy in Lunularia cruciata (L.) Dum.: I. RESUMPTION OF GROWTH AND ITS CONTINUATION

Lunularia cruciata may become dormant at three stages in itslife history: mature thallus, gemma, and spore. The resumptionof growth and its continuation in various conditions have beenstudied in thalli and gemmae. Air-dry, mature thalli of theIsrael strain, planted on a suitable medium, produce adventitiousbranches ventrally from the region immediately posterior tothe existing meristem, which itself fails to resume growth.When dormant gemmae are taken from the gemma-cup, however, theexisting apical cells and meristems simply recommence growing,so that the new growth is continuous with the old. Except inthe case of mature thalli aroused from dormancy, apical dominanceis pronounced, and branching occurs only by bifurcation. Thisdominance can be broken by applying sucrose solution to thegrowing tips—possibly a plasmolytic effect. The growth in area of freshly planted gemmae accelerates forabout 25 days before its relative rate slows to any great extent.During the first half of this period, growth is due exclusivelyto the expansion of existing cells, but subsequently cell numbersincrease rapidly from the 8–10,000 present in the dormantgemma. Cell numbers were estimated by using a modified macerationtechnique, in which chelation followed prolonged fixation. Lunularia grows successfully at quite low light intensities.Of the mineral nutrient deficienccs investigated, lack of Plimits growth most severely, although N-deficiency also restrictsit to a very low level. Gemma-cup production appears to be unaffected by light intensity,at least within levels permitting growth. There is, however,a large temperature effect, cup production decreasing markedlyabove 12° C.     

Evidence for a Flowering Inhibitor Produced in Long Days in Kalanchoe blossfeldiana

Experiments with Kalanchoe blossfeldiana are described in whichperiods of short-day treatment were interrupted by intercalatedlong days or light breaks during long dark periods. The effectsof 24-hour dark periods preceding and following such intercalatedlong days were also investigated. The results of these experiments have shown that: Single longdays intercalated between numbers of short days have a positiveinhibitory effect on flower initiation and are not merely ineffective.The inhibitory effect expressed as the number of inductive cyclesannulled is approximately additive, provided the long days areinterspersed with short days, but not if several long days aregiven consecutively. On the average 1 long day is capable ofannulling the flower-promoting effect of about 1 short days.To a first approximation flower numbers in Kalanchoe increaseexponentially with the number of inductive cycles given—upto at least 12 short days; the inhibitory effect of long daysinterspersed with short days also fits an exponential curve;i.e. the inhibition is roughly proportional to the amount ofprevious photo-periodic induction. A light break of as littleas 30 seconds' duration given in the middle of a long dark periodis as inhibitory as a long day. If followed by a long dark periodthe inhibition of an intercalated long day is almost completelyneutralized; a long dark period preceding it has no such effect. These results have been interpreted as due to the interactionof a flowering inhibitor with a reaction leading to flowering.A mechanism involving competitive inhibition of an adaptivelyformed enzyme has been described as a possible example of thekind of reaction which could account for the results presented.     

Growth and Dormancy in Lunularia cruciata (L.) Dum: VI. GROWTH REGULATION BY DAYLENGTH, BY RED, FAR-RED, AND BLUE LIGHT, AND BY APPLIED GROWTH REGULATORS AND CHELATING AGENTS

Experiments with photoperiods ranging from 2 to 24 h confirmthat 8 h light per day is optimal for Lunularia: there is nogrowth in the dark or in continuous light, which causes therapid onset of dormancy. Short-day cycles intercalated amonga series of continuous light cycles promote growth; in cycleslonger than 24 h very long dark periods are detrimental. Withvery short photoperiods (5 min) red light promotes growth moreeffectively than white light at higher intensity; far-red actsas dark. The growth effects of red and far-red light breaks(3 min) depended on the time of application; red light inhibitedin the middle but promoted at the beginning of the 16-h darkperiod of a short day; far-red light had the opposite effect;in each case red and far-red effects were reversible by theother wavelength. Blue light gave the same response as red includingthe reversibility of far-red effects and vice versa. Surprisingly,significant effects of 5 min red, blue, and far-red irradiationwere also found in the middle of the main high-intensity white-lightperiod, red and blue promoting growth, far-red reducing it;again there was ready reversibility of the effects. Growth promoters of higher plants are generally inhibitory toLunularia or have little effect; among growth retardants TIBA,Phosphon D, and CCC gave a slight promotion of growth. EDTApromoted growth (cell numbers) very significantly while 8-hydroxyquinolinewas initially inhibitory, but had a marked latent promotingeffect when subsequently washed from the thalli.     

Effects of Photoperiod on the Apparent Viscosity of Leaf Cytoplasm in Kalanchoe blossfeldiana: 2. Some Long- and Short-term Effects of Daylength and Spectral Composition of Light

The effects of a range of daylength treatments upon the apparentviscosity of the mesophyll cytoplasm were tested in Kalanchoeand some other daylength-sensitive species by means of a centrifugationtechnique. In Kalanchoe, apparent viscosity increases with increasingshort-day induction, the rise being rapid initially but fallingoff subsequently in the same way as the flowering response.Since the response to light-break treatment is equivalent tolong-day conditions, the effect is truly photoperiodic. Theshort-day increase appears to be only partially reversible uponreturn to long-days. Single leaves may be treated and the effect is not translocatedto the opposite member of a leaf pair. The immediate response to change from light to dark and viceversa is oscillatory; e.g. in the dark an initial fall is succeededby a steep rise and then a more gradual fall in apparent viscosity. Light-breaks with red light are equivalent to white light, andthose with far-red light to darkness. No reversal of red effectsby far-red light has yet been demonstrated. In other species tested, apparent viscosity was significantlyhigher under short-day in Chrysanthemum and Michaelmas Daisy,and under continuous light in Epilobium, i.e. in the daylengthswhich induce flowering. The results are discussed in relationto the present theories of photoperiodism.     

Factors controlling Flowering in the Chrysanthemum: V. DE-VERNALIZATION IN RELATION TO HIGH TEMPERATURE AND LOW LIGHT INTENSITY TREATMENTS

Experiments are described which indicate that the annual vernalizationrequirement of the basal shoots of the Chrysanthemum is dueto annual devernalization of these shoots as the main axis growsup and flowers. Plants sprayed with varying concentrations ofmaleic hydrazide were arrested in their growth for considerableperiods, but this enforced ‘dormancy’ did not affecttheir vernalization status. This makes it appear unlikely thatmere suppression of growth through apical dominance of the mainshoot is the cause of this de vernalization of basal shoots.Fully or partly vernalized plants heated to 40° C. for upto 30 hours did not become dc-vernalized. Heat treatment at35° C. for as long as 30 days also failed to achieve completedc-vernalization, but here flowering was delayed by periodsequivalent to the time spent at high temperature. However, atthe end of the heat treatment progress towards flowering wasresumed at the normal rate. Complete dc-vernalization can bebrought about by prolonged exposure to low intensity illumination.This treatment appears to be effective right up to the stagewhen the first morphological changes leading to inflorescenceformation take place. These results are discussed in relationto similar experiments on the de-vernalization of rye and Hyoscyamusniger.     

Wetting and temperature requirements for infection of mature apples by Venturia inaequalis in South Africa*

Conidia of Venturia inaequalis were used to inoculate mature (picking ripe stage) Granny Smith fruit in greenhouse inoculation chambers and in the orchard. Wetting periods necessary for fruit infection were longer than those previously reported for leaves, while continuous wetting was followed by heavier infection than intermittent wetting. Temperatures during storage affected symptom development and the degree of infection was greatly influenced by the inoculum potential (number of conidia). At high inoculum potentials and with infection indices [fruit wetting period (h) × mean temperature (°C)] of 440–600, 601–1000 and >1000 subsequent disease development was ‘light’, ‘moderate’ and ‘heavy’, respectively if fruit was unprotected by fungicides. It was concluded that after conditions favouring infection, fruit should not be stored for long periods, but should be marketed for consumption immediately after harvest.