The work described in this thesis had several main objects which are summarised as follows: • To study the carbohydrate movement from alga to fungus during photosynthesis in the lichen Xanthoria aureola (Ach.) Erichs. (since this work commenced the variety of Xanthoria parietina used in the experiments has been recognised as a separate species and is referred to by this new name). Previous research on this aspect of the physiology of lichens has been largely limited to Peltigera polydactyla. This lichen contains the blue-green alga Nostoc which produces glucose during photosynthesis that passes to the fungus where it is rapidly converted into mannitol. Xanthoria aureola was selected for the experiments in this investigation because it contains a green alga, Trebouxia, which is the commonest algal symbiont found in temperate lichens. • To study the carbohydrate movement between the symbionts of other lichens to determine the extent to which the pattern found in Peltigera polydactyla and Xanthoria aureola occurred in other species. • To study some general aspects of the carbohydrate metabolism of Xanthoria aureola such as the changes which occur during starvation, the uptake of carbohydrates and the variation in sugar alcohol content at various times of the year. • To study a number of ecological and taxonomic aspects of the biology of Xanthoria parietina sensu lato by transplanting thalli of different varieties between inland and sea shore habitats. 1. Carbohydrate movement from alga to fungus in Xanthoria aureola. a. Products of photosynthesis in the intact thallus. Before investigating the role of the alga and fungus in the symbiosis, it was necessary to study photosynthesis in the complete thallus. When samples of this lichen were floated in the light on aqueous <sup>14</sup>C sodium bicarbonate solutions, <sup>14</sup>C was at first incorporated in substantial amounts into the sugar alcohol ribitol which had not previously been reported in lichens. After several hours <sup>14</sup>C also began to accumulate in mannitol and a small amount was also found in arabitol. It was noted that less than 10% of the fixed <sup>14</sup>C occurred in the ethanol-insoluble fraction over a 24 hour period, and in experiments where the redistribution of a pulse of <sup>14</sup>C-ribitol was studied, it was found that this became largely converted into <sup>14</sup>C-mannitol and <sup>14</sup>C-arabitol over a 48 hour period. b. Studies on the isolated components of the lichen. i. The alga. The alga of Xanthoria aureola, which is a species of Trebouxia, was successfully isolated and maintained in pure culture. Since there was the possibility that the behaviour of the alga in pure culture is different from that in the lichen, a technique was developed for isolating quantities of the alga direct from the thallus. This was done by gently homogenising thallus lobes and subjecting the homogenate to a regime of low speed differential centrifugation so as to produce a moderately pure suspension of living intact algal cells. It was demonstrated that both the cultured and directly isolated alga produced <sup>14</sup>C-ribitol and <sup>14</sup>C-sucrose within the cells during photosynthesis on <sup>14</sup>C-sodium bicarbonate solution. However when the alga had been in pure culture for some time, less <sup>14</sup>C was incorporated into carbohydrates, especially ribitol, than when the alga was directly isolated from the thallus. Both kinds of algal preparation were found to release carbohydrate, mainly in the form of ribitol, into the medium. However they did this only slowly, and the cultured alga appeared to release substances less readily than the directly isolated alga. <strong>ii. The fungus</strong>. This was also isolated into pure culture from fungal ascospores and was subsequently grown in liquid culture. The lichen fungus of Xanthoria aureola was found to grow on a range of sugars and sugar alcohols. It was able to convert <sup>14</sup>C-glucose rapidly into <sup>14</sup>C-mannitol. Under certain cultural conditions it was found also to contain arabitol, confirming that this is a fungal product. c. Attempts to inhibit the movement of carbohydrates between the symbionts If <sup>12</sup>C-pentitols were present in the medium during photosynthesis by Xanthoria aureola on <sup>14</sup>C-sodium bicarbonate solutions, then <sup>14</sup>C-ribitol appeared in the medium and little <sup>14</sup>C-mannitol was formed within the thallus. This was explained in terms of the <sup>14</sup>C-ribitol produced by the alga being unable to compete with the high concentration (l or 2%) of non radioactive pentitols at the fungal uptake sites, so that the entry of <sup>14</sup>C-ribitol into fungus was prevented and it therefore diffused out into the medium. It was only possible to obtain this inhibition with pentitols. Other sugars or sugar alcohols were unable to prevent carbohydrate movement, suggesting that the fungal uptake mechanisms had a high specificity for pentitols. These results suggested that carbon moved between the symbionts principally in the form of ribitol. d. The rate of movement of carbohydrate between the symbionts In Peltigera polydactyla it had been shown by earlier workers that <sup>14</sup>C fixed by the alga moves to the fungal medulla within thirty minutes from the start of photosynthesis on <sup>14</sup>C-sodium bicarbonate. In the type of inhibition experiment described above, at least 25% of the total fixed <sup>14</sup>C was released from samples of Peltigera into the medium within three hours if it contained <sup>12</sup>C-glucose. It was therefore concluded that movement of carbon between the symbionts was rapid. In analogous inhibition experiments carried out here using Xanthoria aureola, it took as long as twenty four hours for a similar proportion of the total fixed <sup>14</sup>C to be released into the medium when it contained <sup>12</sup>C-ribitol, and this suggested that the movement of carbohydrates between the symbionts in this species was much slower. Further experiments were devised in which the rate of transfer of a pulse of <sup>14</sup>C from alga to fungus was estimated both in Peltigera polydactyla and Xanthoria aureola under identical conditions. These confirmed that movement of carbohydrate was much slower in the latter lichen. <strong>2. Studies of carbohydrate movement in other lichens</strong> a. Lichens containing blue-green algae Three lichens were examined, two contained Nostoc (Sticta fuliginosa, Lobaria scrobiculata) and the third, Rivularia (Lichina pygmaea). They all showed similarities with Peltigera polydactyla in that glucose appeared to move between the symbionts and this was converted into mannitol by the fungal part of the thallus. There was evidence that the movement of carbohydrate from alga to fungus was rapid in the species containing Nostoc but slower in Lichina pygmaea. b. Other lichens containing green algae Lichens containing four genera of green algae (including Trebouxia) were studied and it was remarkable that in each case <sup>14</sup>C initially accumulated in sugar alcohols, during photosynthesis. The compounds ribitol, erythritol and sorbitol appeared to be the respective photosynthetic products of the algae Trebouxia (and Coccomyxa), Trentepohlia and Myrmecia.
| Identifer | oai:union.ndltd.org:bl.uk/oai:ethos.bl.uk:644613 |
| Date | January 1967 |
| Creators | Richardson, David H. S. |
| Publisher | University of Oxford |
| Source Sets | Ethos UK |
| Detected Language | English |
| Type | Electronic Thesis or Dissertation |
| Source | http://ora.ox.ac.uk/objects/uuid:7ec5e5e6-a46b-4d1d-8f5a-21293bc0ff29 |
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