Thesis (PhD(Agric)--University of Stellenbosch, 2010. / ENGLISH ABSTRACT: The bright colouration of involucral leaves in Leucadendron is unfortunately transient in nature. Undesirable colour changes render this cut flower unmarketable, resulting in a considerable loss of profit. A deeper understanding of the mechanism leading to colour change is needed to form the framework on which future manipulation strategies can be built. Yellow Leucadendron possess the ability to degreen and regreen naturally, a phenomenon linked to the controlled degradation of chlorophyll and the lesser degradation of carotenoids, which then impart the yellow colour. This colour change is directly linked to the development of the inflorescence. Involucral leaves degreen towards anthesis and are entirely yellow at full bloom. They begin to regreen again when the last florets on the cone have wilted. Deconing before flowering completely inhibits the colour change. Deconing at full bloom, results in leaves regreening sooner. Therefore the inflorescence appears to be the origin of the cue for colour change. Any factor that expedites the death of the florets, results in sooner regreening of involucral leaves. Ultra-structurally, the degreening and regreening resulted from a transdifferentiation of mature chloroplasts to gerontoplast-like plastids, which upon regreening completely redifferentiated into fully functional chloroplasts. In the red Leucadendron cultivar Safari Sunset, the photosynthetic pigment degradation pattern is identical to that of yellow cultivars. However, colour expression is complexed by the presence of anthocyanins. Anthocyanin concentration was shown to be directly related to the opening of the flower head rather than to the phenological development of the inflorescence. With opening, the previously shaded inner involucral leaf surfaces are exposed to higher levels of irradiance and respond by turning red, presumably for photoprotection. Similar to yellow cultivars, any factor leading to the death of the florets before flowering, not only prevents the degreening of involucral leaves, but also prevents the opening of the flower head and therefore the associated change in anthocyanin levels. The ecological significance of regreening was also investigated. What does a female Leucadendron plant stand to gain by regreening rather than discarding the involucral leaves? Regreened involucral leaves were shown not to play a significant role in providing photosynthates for the developing cone. Although the presence of regreened involucral leaves were shown to provide protection against high irradiance and radiant heating of the cone, they were not essential to ensure survival of the cone. The small floral bracts were shown to be very capable of adaptation. The most plausible reason for regreening is therefore assumed to be based on a cost-benefit relationship. As most Leucadendron are adapted to grow on very nutrient poor soils, the question should maybe be rephrased. Why waste valuable resources? Sclerophyllous leaves, like the involucral leaves, are costly to make and therefore reusing, rather than discarding them does seem a sensible strategy for survival. / AFRIKAANSE OPSOMMING: Leucadendron snyblomme word gekenmerk deur die helder kleure van hul omwindselblare. Die helder kleure is egter slegs vir 'n kort periode aanwesig waarna die snyblomme onbemarkbaar word, met aansienlike verlies aan potensiele inkomste. Die ontwikkeling van manipulasies ten einde die bemarkbare periode van Leucadendron te verleng, berus op die verkryging van 'n dieper insig in die meganisme van kleurverandering. Die kleurveranderinge van geel Leucadendron omwindselblare is te wyte aan 'n unieke vermoë tot die gereguleerde degradasie en heropbou van chlorofiele en karotenoiede onder direkte beheer van die ontwikkelende bloeiwyse. Met die aanvang van blom, lei groter proporsionele degradasie van chlorofiele tot geleidelike vergeling van omwindselblare. Die hele blomhofie verkry uiteindelik met volblom 'n helder geel kleur. Sodra die laaste blommetjies doodgaan, neem chlorofiel- en karotenoiedsintese weer in aanvang en binnekort is die omwindselblare weer net so groen soos voor die aanvang van blom. Die geel verkleuring kan verhoed word deur die keël voor blom uit te breek. Enige faktor wat die dood van die blommetjies versnel, asook die uitbreek van keël tydens volblom, lei tot die vroeëre aanvang van vergroening. Die degradasie van plastiedpigmente hang nou saam met die differensiasie van volwasse chloroplaste tot gerontoplast-agtige plastiede wat op hul beurt weer tydens vergroening tot volkome funksionele chloroplaste herdifferensieer. Soortgelyk aan geel Leucadendron kultivars, vind die veranderinge in plastiedpigmente ook plaas tydens blom van die rooi kultivar, Safari Sunset. Kleurveranderinge in 'Safari Sunset' is egter meer ingewikkeld vanweë die aanwesigheid van variërende konsentrasies antosianiene. Antosianienkonsentrasies en rooi kleur neem toe tydens blom vanwee die blootstelling van die beskutte adaksiale binnekante van omwindselblare aan hoe irradiasie met die oopvou van die blomhofie. Die akkumulasie van antosianiene het moontlik 'n fotobeskermende funksie. Kleurveranderinge in 'Safari Sunset' kan, soos in geel kultivars, voorkom word deur blom te verhoed. Antosianiensintese word voorkom deurdat die blomhofie geslote bly en is nie direk gekoppel aan blom soos wat met plastiedpigmente die geval is nie. Die belang van vergroening is ondersoek na aanleiding van die vraag oor wat dit 'n vroulike Leucadendron baat om omwindselblare te behou na die afloop van blom? Die bydrae van foto-assimilasie deur omwindselblare tot die ontwikkeling van keels is beperk. Alhoewel omwindselblare wel keels teen hoe irradiasie en stralingsverhitting beskerm, is die blomskutblare in staat om aan te pas by hierdie kondisies. Die mees waarskynlike verklaring vir die behoud van die omwindselblare na blom berus moontlik op 'n koste-voordele verwantskap. Alhoewel nie essensieel nie, is die beperkte bydrae van die omwindselblare na die afloop van blom tot die oorlewing en welstand van die keel waarskynlik genoegsaam om hul behoud te regverdig. Verskeie Leucadendron spesies groei in gronde wat baie arm is aan nutriente. Sklerefiele blare, soos die van Leucadendron, is verder duur om te vervaardig. Dit maak dus sin om hulle vir meer as een funksie te herontplooi eerder as om hulpbronne te belê in meer gespesialiseerde en minder durende blombykomstighede. Dus dui die behoud van omwindselblare dalk op 'n strategie wat gemik is op die behoud en besparing van beperkte hulpbronne.

Thesis (MScAgric (Horticulture))--University of Stellenbosch, 2010. / ENGLISH ABSTRACT: Peel colour is an important quality factor in the production of bi-coloured apple fruit. Most markets set minimum requirements for red colour coverage. Fruit that do not meet these requirements are downgraded and has a major impact on the profitability of apple production in South Africa. South African apple production areas are amongst the warmest in the world. Since anthocyanin accumulation requires induction at low temperature and synthesis require mild temperatures, experiments were conducted to investigate optimum day and night temperatures for red colour development throughout fruit development for red and bi-coloured apple cultivars grown in South Africa. We found that redder strains of bi-coloured apple cultivars did not appear to owe their enhanced pigmentation to higher temperature optima for anthocyanin synthesis. The optimum day temperatures for red colour development in the different cultivars seemed to differ between seasons, but not between production areas. In general, red colour in the cultivars evaluated developed maximally between 17 ºC and 25 ºC. The optimum day temperature for red colour development remained constant throughout fruit development for most cultivars, but increased roughly from 14 ºC to 22 ºC in ‘Cripps’ Pink’ between January and April. The extent of red colour development increased during fruit development in all the cultivars assessed. We were unable to determine optimum induction temperatures for red colour development. ‘Royal Gala’ from Ceres seemed to benefit from induction at 4 ºC while red colour in ‘Fuji’ decreased with decreasing temperature. To explain the presence of anthocyanins in immature apple fruit, we tested the hypothesis that anthocyanins protect the peel from photoinhibition and photooxidative damage during conditions of increased light stress. First we established that the rate of colour change in response to a passing cold front appears to be sufficient to provide photoprotection during a cold snap. Also in agreement with the hypothesis, ‘Cripps Pink’ peel incurred significantly more photoinhibition at low temperature (16 ºC) compared to mild (24 and 32 ºC) and high (40 ºC) temperature under high irradiance with visible light. Recovery rate was temperaturedependent, being the slowest at low temperature and increasing with temperature. The photoapparatus in ‘Cripps Pink’ peel appears to be particularly sensitive to light stress at low temperature throughout the season, with significant photoinhibition occurring even at moderate temperature (24 ºC). The sensitivity of the apple peel to photoinhibition increased throughout the season at lower irradiance levels, but remained the same at higher irradiance. In our final experiment, fruit were exposed to high irradiance at low and mild temperature before exposure to high temperature in combination with high irradiance. This was done to test the hypothesis that photoinhibition incurred during cold snaps predisposes peel to photothermal damage when temperature increases again after the cold snap. Unfortunately, due to the severity of the stress incurred in response to high temperature treatment, the results were inconclusive. / AFRIKAANSE OPSOMMING: Vrugkleur is ‘n belangrike kwaliteitsfaktor in die produksie van tweekleurappels. Die meeste markte stel minimum vereistes vir rooi kleurbedekking. Vrugte wat nie aan hierdie vereistes voldoen nie, word afgegradeer. Suid-Afrika se appel produksie areas word beskou as van die warmste ter wêreld. Antosianien akkumulasie benodig induksie by lae temperature gevolg deur sintese in lig by matige temperature. Gevolglik het swak rooi kleurontwikkeling onder plaaslike toestande ‘n groot impak op die winsgewendheid van appelproduksie in Suid-Afrika. Eksperimente is uitgevoer om die optimum dag- en nagtemperature vir rooi kleurontwikkeling tydens vrugontwikkeling vir die rooi en tweekleur appel kultivars wat in Suid-Afrika geproduseer word te bepaal. Ons het gevind dat die verhoogde pigmentasie van rooier seleksies van tweekleurappel kultivars nie aan ‘n hoër temperatuur optimum vir antosianiensintese toegeskryf kan word nie. Die optimum dag temperature vir rooi kleurontwikkeling vir die onderskeie kultivars verskil klaarblyklik tussen seisoene, maar nie tussen produksie areas nie. Oor die algemeen het kleurontwikkeling maksimaal plaasgevind tussen 17 ºC en 25 ºC. Die optimum dagtemperatuur vir rooi kleurontwikkeling het konstant gebly tydens vrugontwikkeling, buiten vir ‘Cripps’ Pink’ waar dit toegeneem het van ongeveer 14 ºC tot 22 ºC vanaf Januarie tot April. Die mate van rooi kleurontwikkeling het in al die kultivars toegeneem deur die loop van vrugontwikkeling . Ons kon nie daarin slaag om optimum induksie temperature vir rooi kleurontwikkeling vas te stel nie. Rooi kleurontwikkeling van ‘Royal Gala’ uit Ceres is moontlik bevorder deur induksie by 4 ºC, terwyl ‘Fuji’ se rooi kleur afgeneem het met ‘n verlaging in induksie temperatuur. Ten einde die teenwoordigheid van antosianien in onvolwasse appelvruggies te verduidelik, het ons die hipotese getoets dat antosianien die vrugskil beskerm teen fotoinhibisie en fotooksidatiewe beskadiging gedurende tydperke van verhoogde ligstres. Eerstens het ons bevestig dat die tempo van kleurontwikkeling in reaksie op ‘n koue front waarskynlik vinnig genoeg is om fotobeskerming te verleen. Vervolgens is gevind dat ‘Cripps’ Pink’ vrugskil aansienlik meer fotoinhibisie ervaar het by lae temperatuur (16 ºC) in vergelyking met matige (24 ºC en 32 ºC) en hoë (40 ºC) temperatuur onder hoë irradiasie met sigbare lig. Die hersteltempo was temperatuur-afhanklik; dit was die stadigste by lae temperatuur en het toegeneem met ‘n toename in temperatuur. Die foto-apparaat in ‘Cripps’ Pink’ vrugskil blyk besonder sensitief te wees vir ligstres by lae temperatuur regdeur die groeiseisoen met aansienlike fotoinhibisie by selfs matige temperatuur (24 ºC). Die sensitiwiteit van die vrugskil vir fotoinhibisie het toegeneem deur die groeiseisoen by laer ligvlakke, maar het dieselfde gebly by hoër vlakke van irradiasie. Laastens is vrugte blootgestel aan hoë irradiasie by lae en matige temperatuur voordat dit vervolgens blootgestel is aan hoë temperatuur in kombinasie met hoë irradiasie. Dit was om die hipotese te toets dat fotoinhibisie wat opgedoen word gedurende ‘n onverwagte koue periode, die skil meer vatbaar maak vir fototermiese skade sodra die temperatuur weer styg na die koue periode verby is. Ongelukkig het die hoë temperatuur stres al die behandelings tot so ‘n mate geaffekteer dat dit onmoontlik was om enige gevolgtrekkings vanuit ons resultate te maak.