Volume 16(1), 212-215, 2012 JOURNAL of Horticulture, Forestry and Biotechnology www.journal-hfb.usab-tm.ro Aspects regarding the physiology of cuscuta (Cuscuta L. Convolvulaceae) Tănase Maria1*, Sand Camelia1 1 “Lucian Blaga” University of Sibiu *Corresponding author. Email: [email protected] Abstract Many species of Cuscuta contain a small amount of chlorophyll; and while both chlorophyll a and b are present in normal proportions, they only occur in a very low concentration. Several authors have demonstrated 14 the capacity of Cuscuta for photosynthesis, making use of CO2 . Fer has demonstrated that Cuscuta exposed to strong air illumination, with a CO 2 concentration much higher than the normal atmospheric CO 2 concentration, fixes more CO2 during the photosynthesis process than the amount produced by means of respiration in both independently living seedlings and in the stem excised from the host. The migration of assimilates is only occurring acropetally, not only in the case of the plantula, but also in the case of the mature parasite. The entirely acropetal migration of assimilates can be explained by the attracting effect exerted by the terminal sprout, as well as by the lateral sprouts situated on the subterminal part of the branch. The rudimentary root of the plantula does not have a meristem [19]; [12]; [28]; [23]; and the absence of a downward migration within the plantula thus appearing to be determined by the absence of the radicular meristem [9]. Photosynthetic apparatus and chlorophyll degree of concentration. With the exception of its roots, the organs of Cuscuta have only a low grade of chlorophyll concentration, and are ‘equipped’ with stomates [19];[29];[4]. According to several studies carried out by means of an electronic microscope, the chloroplasts of several Cuscuta species have a rudimentary structure and contain substantial amounts of starch granulates. The occurrence of the chlorophyll a and b in numerous Cuscuta species is certain. The results of MacLeod [17] and Laudi [15] indicate the chlorophyll grade of concentration of these parasites; a smaller one than that of autotrophic plants. With the exception of Laudi [15], all authors agree that the value of the proportion chlorophyll a / chlorophyll b is of between 2.2 and 3.5; a value which is analogous to that occurring with most autotrophic plants. Highlighting and measuring the intensity of photosynthesis. One has noted that the plantulae situated under light/brightness conditions liberate oxygen [8];[24] and the quantity of oxygen is directly proportional to the lighting of the plantulae. Moreover, the CO2 anchorage and its stimulation by brightness have been noted as well [16] and [18]; [20]; [1] and [2]; [14]; [13]. Among the authors who acknowledged the occurrence of a photosynthetic CO2 anchorage when it comes to Cuscuta, Ismail and Obeid [13] attempted to establish an accurate balance of the 212 Key words Cuscuta sp., photosynthesis, chlorophyll a and b, assimilate transport phenomenon, based on measuring the intensity of the photosynthesis, resp. of the respiration. According to their results, when it comes to the Cuscuta hyalina, in an atmosphere which is not enriched with CO2, the intensity of photosynthesis and that of respiration appear to be equal. In this context one has to raise the following question: to what degree is the photosynthesis capable of allowing the development of the plantula Fer’s experiments [9] confirmed Pizzolongo’s results [22], showing that in an atmosphere which is not enriched with CO2, the photosynthesis cannot compensate for the respiration. Under these conditions, the growing of the plant is not possible, and the intensity of respiration, resp. photosynthesis thus explains the species’ parasitism. Therefore, it becomes obvious that – since its growth is fast – the plant exhausts the resources contained in its seed rapidly, thus dying unless it anchors on a host plant. The photosynthesis is not intense enough in order to allow the growth of the parasite by itself; the latter thus needing a host plant to provide itself with carbonated substances. If the atmosphere has a higher degree of CO2, the photosynthetic intensity of Cuscuta will be increased. Under such conditions, despite its low degree of chlorophyll, Cuscuta is able to provide – if not all, then at least a significant part of those substances which are necessary for its growth. Table 1 Bibliographic data regarding the chlorophyll degree of concentration in Cuscuta Degree of Culture conditions Chlorophyll development / Cl a / Authors ( g/g of fresh host plant Cl b matter) C. reflexa Parasite, on natural 70 (1*) 2,8 Mac LEOD [17] Nicotiana C.campestris Parasite, on natural 37 2,2 Mac LEOD [17] Bulbine C. pentagona Plantulae, 4-7 Continuous - ( 2*) 2,7 Zimmerman [29] days lighting (2000 Lux) C australis Plantulae, 4-7 Continuous - ( 2*) 2,7 Bertossi et al. [3] days lighting (1000 Lux) C. epithymum plantulae Not mentioned - ( 2*) 3,3 - 3,5 Sala et al. [24] C. australis parasite (host natural - ( 2*) 3,5 Baccarini et al. [2] not mentioned ) C. australis parasite (host natural 145 (3*) 1 Laudi [15], Tanase [27] not mentioned ) (1*)- if the host plant, exhausted by the parasite, any comparison between haustoria and roots; a is dying, Cuscuta reflexa, ‘greened’ by the high degree comparison to which several authors tended to refer in of chlorophyll, reaches its highest values; their works (Pierce, [21] and Fratianne, [11)]. (2*)- the studied data does not allow the In the case of Cuscuta, once the vascular calculation of chlorophyll concentration; connections between parasite and host plant have been (3*)- gall stems (produced by the Cuscuta established, haustoria no longer develop [26]; in ladybug – Smicronyx sp.) have a very high degree of conclusion, in a functional haustorium no single spot chlorophyll concentration. characterized by a merismatic occurrence persists. Thus it can be explained why the Cuscuta haustoria do not exert any attracting effect, and why no downward The assimilates’ migration within the plantula. A significant part of assimilates synthesized migration of assimilates occurs. by the plantula migrate toward the stem’s apex. This point of view is confirmed by the results of Fer’s Material and Method experiments [10], according to which, there is no downward migration of assimilates, the only occurring Several Cuscuta plants have been used, both migration being acropetal, not only in the case of the independent and parasitic ones, in order to highlight plantula, but also in that of the mature parasitic plant. and measure the photosynthesis intensity under certain The rudimentary root of the plantula lacks a conditions; not only normal atmospheric ones, but also meristem character [19]; [12]; [29];[23], whereas the carbon dioxide-enriched atmospheric conditions. absence of any downward migration within the The migration of assimilates – which has shown plantula seems to be determined by the absence of the only an upward tendency - has also been analyzed; the radicular meristem [9]; [31]. Even if it plays a certain lack of a downward migration has been highlighted by role with regard to the absorption of mineral elements, samples of five-days-aged plantulae. the rudimentary root of the plantula does not represent any attraction for the assimilates, due to the meristem’s Obtained results absence. The lack of a downward migration explains the limited duration of the root, which exhausts its According to the results of our research, the resources and dies unless it is provided with migration of assimilates is only occurring acropetally, assimilates. not only in the case of the plantula, but also in the case Similarly, when it comes to the parasite plant, of the mature parasite. This result can be accompanied haustoria do not represent any centre of attraction for by several remarks: the assimilates, even if they ensure the absorption of Downward migration can be induced on the water and dissolved substances. The attracting effect parasital plant by leaving a single sprout at the basis of exerted on the assimilates by the roots of the superior the branch. The entirely acropetal migration of autotrophic plants [5];[25];[6];[7] occurs neither at the assimilates can also be explained by the attracting rudimentary root of the plantula, nor at the adult plant effect exerted by the terminal sprout, as well as by the haustoria. This constitutes a decisive argument against Analyzed species 213 lateral sprouts situated on the subterminal part of the branch. When it comes to the plantula, the only occurring vegetative spots are the terminal sprout as well as two or three lateral sprouts, situated quite close to the terminal one. The rudimentary root of the plantula does not have a meristem [19]; [12]; [28]; [23]; the absence of a downward migration within the plantula thus appearing to be determined by the absence of the radicular meristem [9]. For the sake of comparison, it would be interesting to present some findings regarding the migration of assimilates in the case of other parasitic fanerogams, namely: hemiparasites on the root (Scrophulariaceae) and hemiparasites on the stem (Loranthaceae). On the studied Scrophulariaceae, the assimilates synthesized by a branch migrate to different parts of the parasite. One has to note the basipetal migration of the synthesized part towards the root of the parasite, as it is the case with the Striga senegalensis, parasite on the Sorghum vulgare and with the Odontites verna, parasite on the Hordeum vulgare[10] . Concerning the hemiparasital Loranthaceae, one can mention two different cases: - The first, most often-occuring case refers to a situation in which marked assimilates, synthesized by a branch of the parasite, migrate to all directions, including towards haustoria, but without passing into the phloem of the host plant. This is the case of the Viscum album, parasite on the Populus nigra [10], of the Phoradendron sp, parasite on several gymnosperms as well as a large number of wood dicotyledon as well as Tapinanthus bangwensis, parasite on the Theobroma cacao [10]. - The second case, still controversial, is that of the Arceuthobium sp, parasite especially on the species Abies and Pinus. According to Redisske and Shea, a part of assimilates synthesized in the parasite’s branches migrate towards the haustoria, thus in a basipetal direction [10]. Conclusions Thus, among the studied parasitic fanerogams, the lack of downward migration, which has been highlighted in the case of Cuscuta, seems to occur only in the case of the Arceuthobium species. In order to explain the occurrence of a downward migration in the case of parasitic fanerogams, one could formulate the following hypothesis: Regarding the hemiparasites belonging to the Scrophulariaceae family, only the extremities of certain roots come into contact with the roots of the host plant, thus forming haustoria . Therefore, there are free roots whose apex can exert an attracting effect on the assimilates. Similarly, in the cases of the Loranthaceae family, the Viscum, Phoradendron and Arceuthobium genera, beside primary haustoria there are cortical cords, of indefinite 214 growth, which spread around the primary haustoria. It therefore seems logical to think that the extremities of these cords have a merismatic character, thus constituting centres of attraction for the assimilates. In the case of the Arceuthobium genus, the lack of downward migration is justified by the absence of the differentiated phloem [10]. When it comes to Cuscuta, once the vascular connections between parasite and host plant have been established, haustoria no longer develop [26]. In conclusion, in the case of a functional haustorium no single zone characterized by a merismatic occurrence persists. Thus it can be explained why the Cuscuta haustoria do not exert any attracting effect and why no downward migration of assimilates occurs. The lack of a sampled downward migration from the host to the Cuscuta lupuliformis partly explains the deterioration of the older branches of the parasite. In fact, it has already been shown that a potential Cuscuta haustorium does not have any spot preserving a merismatic character. This finding can be explained by the fact that during their ageing process, absorbing cells progressively lose their ability of sampling metabolites. Regarding the assumption of metabolites, a Cuscuta haustorium is functional only for a very limited time. At the end of this phase, the branch situated above will no longer be nourished with metabolites, thus starting to deteriorate. A further assumption explaining the deterioration of the older part of the parasite is the following one: those leaves of the host plant which are parasitized by a haustorium for several weeks in a row, display obvious signs of a progressive deterioration (peripheral yellowing of the foliar limb, then desiccation), which then disperse in a centripetal way, and which eventually has an effect on the entire limb as well as on that part of the petiole situated over the haustorium. From that moment the quantity of metabolites sampled by such a haustorium must be extremely reduced or even null. The deterioration of a Cuscuta branch occurs when it is no longer nourished by the upward flux of the sampled assimilates in the host. The deterioration of the branches is accelerated when they have fruit. Acknowledgement This work was cofinanced from the European Social Fund through Sectoral Operational Programme Human Resources Development 2007-2013, project number POSDRU/89/1.5/S/63258 ”Postdoctoral school for zootechnical biodiversity and food biotehnology based on the eco-economy and the bio-economy required by eco-san-genesys”. Bibliografy 1.Baccarini A., - 1966, Autotrophic incorporation of 14 CO2 in Cuscuta australis in relation to its parasitism. Experientia, 22 2. Baccarini A., - 1967, Doder's autotrophy in parasiting state. Z. Pflanzenphysiol., 57 3.Bertossi F., Baccarini A. si Bagni N., - 1964, Rapporto tra le clorofille a et b in Cuscuta australis. Giorn. Bot. Ital., 71 4. Bhattacharyya P.K., - 1973, Stomata of the section Collienche (Cuscuta ceae). Sci. and Cult., India, 39 5.Bonnemain J.L.,.- 1972, La plante supérieure, siége d’une circulation complexe. Physiol. Vég., 10 6.Couillault J. si Bonnemain J. 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