On the improved vigour of the root-knot nematode infected cowpea plant with phenols of palmyrah wood sawdust extract, fortified with gibberellic acid and indoleacetic acid Proceedings: Animal Sciences, Aji Balaji. A short summary of this paper.
On the improved vigour of the root-knot nematode infected cowpea plant with phenols of palmyrah wood sawdust extract, fortified with gibberellic acid and indoleacetic acid. Indian Acad. I, January , pp. With NPK solution as the basic nutrient applied to the root-knot nematode infected cowpea plants, amendments were made with ppm of phenol prepared from alcoholic extract of palmyrah wood sawdust. Amended nutrient applications reduced galling, fecundity of the pathogen, pathogenic impact, improved root weights and synthesis of metabolites which signified the improved vigour of the infected host.
Meloidogyne incognita; Vigna unguiculata; sawdust extract; gibberellic acid; indole acetic acid; vigour. Introduction While the various enzymes in plant parasitic nematodes account for the tissue lysis Roy , , , many of the breakdown products like phenols, uronic acid com- plexes and disaccharides exhibit compatibility with the host plant tissue metabolites and also with the nematode's enzymes, resulting in altered substrate relations and catalytic repression of the nematode's enzyme Feldman and Hanks ; Giebel , ; McIntyre ; Wallace , such that the feeding and subsequent progeny output of nematode are lowered, with consequent lesser pathogenic impact.
This is finally reflected as tolerance in the susceptible plants. These observations led to the reckoning of phenols, proteins, etc..
At the same time accumulation of indole acetic acid, IAA indole compounds Balasubramanian and Rangaswami ; Setty and Wheeler had also been suspected to be one of the possible reasons for galling in root-knot nematode infections. The higher level of IAA in the galled zones and the normal tissue differentiation adjacent to the galls and the varied influences of IAA and gibberellic acid GA in plant pathogenesis Goodman et al reflect the active IAA-GA relations in the pathological meta- bolism.
While breeding for nematode resistant crops is a welcome feature, resistance breaking biotypes of nematode do also evolve and hence the problem goes unabated.
Singh and Sitaramaiah , stated that the nemastatic factors attributable to sawdust as organic amendment were due to the phenolics released from sawdust. In our laboratory, free phenol in varying dilutions in water, inhibited the hatching of Meloidogyne incognita eggs.
Further, in one susceptible host cowpea , whereas 1: dilution did not appreciably retard the pathogenic impact, and whereas I : 10 retarded the plant growth, 1: dilution yielding ppm of phenol checked the infection, without affecting the host plant.. In view of the last observation, experiments were conducted to assess the efficacy of phenols from the alcohol extract of palmyrah wood sawdust, as well as the influence of hormone and auxin, viz.
GA and IAA on the root-knot nematode pathogenesis in cowpea. Thirty pots were filled with sand. Large increases in the amount of phenolic compounds can occur in stressed plants and those undergoing mechanical damage.
Plant phenolics are considered to have a key role as defence compounds when environmental stresses such as bright light, low temperatures, pathogen infection, herbivores and nutrient deficiency can lead to increased production of free radicals and other oxidative species in plants.
A growing body of evidence suggests that plants respond to these biotic and abiotic stress factors by increasing their capacity to scavenge reactive oxygen species.
In addition, in order to establish a protective role for a given metabolite, it is necessary to monitor concentrations over the life cycle of the plant, to survey plant populations, to determine specific localisation within tissues and to carry out bioassays against insects and microorganisms. Finally, changes in secondary chemistry may also occur during ontogeny and protection may be restricted to the most vulnerable plant organs Robbins et al.
Some soluble phenolics, for example chlorogenic acid, are widely distributed, but the distribution of many other structures is restricted to specific genera or families making them convenient biomarkers for taxonomic studies.
Even if the potential value of plant sec- ondary metabolites to taxonomy has been recognised for nearly years, their practical application has been restricted to the twentieth century and predominantly to the last 40 years. The use of secondary compounds has clear advantages over the use of primary com- pounds in establishing phylogenetic relationships because differences in the complement of secondary compounds are qualitative differences whereas differences in the concen- trations of primary compounds are quantitative differences, and these are subject to both environmental and genetic control.
Phenolic compounds are often similar within members of a clade and therefore the existence of a common pattern of secondary compounds may indeed provide much clearer evidence of common ancestry than morphological similarities attributable either to common ancestry or to convergent evolution Bell, ; Lattanzio et al. The precise way in which plants adapt to low temperature is obviously of scientific interest, but there are also practical and economic aspects. Several studies have suggested that exposure to low temperatures usually triggers a variety of biochemical, physiological and molecular changes that allow the plants to adjust to stress conditions and this response is characterised by a greater ability to resist injury or survive an otherwise lethal low temperature stress.
Lowering temperatures will thermodynamically reduce the kinetics of metabolic reactions. Exposure to low tem- peratures will shift the thermodynamic equilibrium so that there is an increased likelihood of non-polar side chains of proteins becoming exposed to the aqueous medium of the cell. This leads to a disturbance in the stability of proteins, or protein complexes and also to a disturbance of the metabolic regulations.
Lower temperatures induce rigidification of mem- branes, leading to a disturbance of all membrane properties permeability, electric field, cation concentration and water ordering, and this leads to disturbance of the conformation and thus the activity, of membrane-bound enzymes. Chilling is also associated with the ac- cumulation of reactive oxygen species ROS. The activities of the scavenging enzymes will be lowered by low temperatures, and the scavenging systems will then be unable to counter- balance the ROS formation that is always associated with mitochondrial and chloroplastic electron transfer reactions.
The accumulation of ROS has deleterious effects, especially on membranes. Some plants are able to adapt through mechanisms based on protein synthe- sis, membrane composition changes, and activation of active oxygen scavenging systems. Low temperature stress induces accumulation of phenolic compounds that protect chilled tissues from damage by free radical-induced oxidative stress. Many papers report the effects of low temperature on phenolic metabolism, and these have shown that phenolic metabolism is enhanced under chill stress and that the behaviour of the same metabolism is further dependent on the storage temperature.
There is a low critical temperature below which an increase of phenylpropanoid metabolism is stimulated during the storage of plant tissues and this temperature varies from com- modity to commodity.
Figure 1. During the first 60 days of cold storage, there is a relevant increase in flavonoid content, but flavonoid content gradually decreases in fruits stored for a longer period. The timing of the observed peak in the phenol level during cold storage depends on the species or cultivar, the harvesting time and the storage conditions Lattanzio et al.
Generally, this low temperature effect on the phenol level involves a cold-induced stimulation of PAL, the branch point enzyme between primary shikimate pathway and secondary phenolic metabolism.
The observed increases in PAL activity induced by low temperature might involve both enzyme de novo synthesis and release of PAL from a pre-existing but inactive enzyme—inhibitor complex. It is likely that endogenous ethylene, produced in plant tissue exposed to low temperature stress, promotes the induction of PAL activity and this is consistent with data showing that cold-induced PAL activity is reduced by inhibitors of ethylene production or by inhibitors of the action of ethylene.
The onset of ethylene production in stressed plant tissues occurs at approximately the same time as an increase in PAL activity. Moreover, the effect of exogenously-added ethylene on most tissues is to cause increased production of PAL. Low temperature induction of PAL activity alone in plant tissues does not produce a corresponding increase in phenol production.
At low temperatures, it is possible that the subsequent steps in the biosynthesis of phenolic compounds may limit their formation. In this connection, reference must be made to some excellent papers showing that other enzymes important in the phenolic biosynthetic pathway e. This phenomenon is largely dependent on the plant material studied, the storage temperature and the controlled or modified atmosphere used.
Low temperature stress, besides affecting enzymes involved in the general phenylpropanoid pathway, also affects CHS the key enzyme of the flavonoid pathway. The increase in these enzymes of phenolic metabolism presumably contributes to the increased production of phenols at low temperature. An increase in the activity of the enzymes, as well as in the level of phenolic compounds, could combine with the temperature-dependent phase changes in the cellular membrane, to affect the shelf life of stored fruit and vegetables by providing an adequate substrate to the browning reactions.
Browning in plant tissues during handling and storage of fresh fruit and vegetables commonly result from either non-enzymatic or enzymatic reactions involving plant phenols, oxygen and environmental contaminants such as metal ions. Enzymatic browning in fruit and some vegetables starts with the enzymatic oxidation of phenols by polyphenol oxidases PPOs, EC 1. The quinone products can then polymerise and react with amino acid groups of cellular proteins, resulting in black or brown pigment deposits melanins.
Such damage causes considerable economic and nutritional loss in the commercial production of fruit and vegetables. PPOs are located in plastids, and they are not integral membrane proteins, although they are membrane associated. Non-enzymatic causes of browning in plant tissues may be attributable to the interactions between phenols and heavy metals — especially iron — which yield coloured complexes. It is generally accepted that a dark coloured complex of ferric iron and an orthodihydric phenol is responsible for discolouration.
It has been suggested that a phenolic compound involved may be chlorogenic acid 5-O-caffeoylquinic acid and that subcellular decompartmentalisation of plant cells during senescence allows the organic ligand to chelate the iron. Since the metal is originally present in the reduced state, a colourless complex is first formed and when exposed to oxygen, oxidises to yield a coloured compound. Plate 1. It is noticeable that in dis- coloured tissues, phenol content is higher than in the healthy tissues of the same artichoke bract, which does not agree with the hypothesis of enzymatic browning.
In this case we would expect a remarkable lowering of phenol content, due to the enzymatic oxidative phenomena. Furthermore, when artichoke tissues suffered enzymatic browning after me- chanical damages and brief exposure to air, the phenolic content found was much lower than that of intact tissues.
It has been suggested that at pH 6. In plants, ferritin is known to be present in chloroplasts, and, especially, in the plastids of non-photosynthesising tissues.
Several chelating agents are able to promote the release of ferritin iron in the presence of a reducing agent. It has been shown that plant phenols, including caffeic acid and chlorogenic acid, can promote the reductive release of ferritin iron: a direct correlation exists between oxidation—reduction potential and the rate of iron release.
In addition, reductant access to the ferritin iron core is also likely, when molecules are relatively small, Price, ; Boyer et al. On the other hand, PPO activity did not change significantly during the cold storage period. The increased content of phenolics provided an adequate substrate for the browning. These reactions started from the chloroplasts, considered to be the site of chlorogenic acid biosynthesis Ranjeva et al.
This complexed phenolic substrate, removed from the regular post-harvest metabolism occurring during cold storage of ar- tichoke, was released in the free form when acidic pH conditions of the medium during HPLC analyses of artichoke caffeoylquinic acids caused the complex to break down. Ehrlich and Raven were among the first to propose a defined ecological role for plant secondary metabolites as defence agents against herbivorous insects.
These substances are repellent to most insects and may often be decisive in patterns of food plant selection. Through occasional mutations and recombination, angiosperms have produced a series of chemical compounds not directly related to their basic metabolic pathways, but not inimical to normal growth and development.
Most research concerning insect anti-feeding agents has shown the involvement of phenylpropanoids, flavonoids and lignans in the plant resistance mechanism against insects.
The concentration of the phenolic compounds in the plant is a key factor in deterrence and it is the accumulation of phenols in particular parts of the plant that represents a feeding barrier. The effectiveness of phenolics as a resistance factor to animal feeding is enhanced, as aforesaid, by oxidation to polymers, which reduces digestibility, palatability and nutritional value Ananthakrishnan, ; Lattanzio et al.
Plants encounter numerous pests and pathogens in the natural environment. An appro- priate response to attack by such organisms can lead to tolerance or resistance mechanisms that enable the plant to survive Paul et al.
Most plants produce a broad range of secondary metabolites that are toxic to pathogens and herbivores, either as part of their normal programme of growth and development or in response to biotic stress Treutter, ; Agati et al. Both tolerance and re- sistance traits require the reallocation of host resources, therefore defensive chemicals are considered to be costly for plants, reducing the fitness of the host in the absence of disease, because resistance genes might impose metabolic costs on plants e.
One way for a plant to reduce these costs is to synthesise defence compounds only after there has been some degree of initial damage by a pathogen or insect: this strategy is inherently risky because the initial attack may be too rapid or too severe for an effective defence response. Fortunately for plants, their relationship with fungi is usually a mutually beneficial one saprophytic fungi, mycorrhizae and endophytes. A small minority of fungal species has developed further and broken the fine balance of mutual benefit to become plant pathogens.
The first demonstrated example from the early plant pathology literature of phenolic compounds providing disease resistance was the case of coloured onion scales accumulating sufficient quantities of catechol and protocatechuic acid to pre- vent the germination of Colletotrichum circinans spores Link et al. Pre-formed antibiotic phenolics phytoanticipins are stored in plant cells mainly as inactive bound forms but are readily converted into biologically active antibiotics by plant hydrolysing enzymes glycosidases in response to pathogen attack.
These compounds are considered as pre-formed antibiotics because the plant enzymes that activate them are already present but are separated from their substrates by compartmentalisation, enabling rapid activation without a requirement for the transcription of new gene products Osbourn, ; Lattanzio et al. When a pathogen manages to overcome constitutive defence barriers, it may be recognised at the plasma membrane of plant cells.
Activation of inducible plant defence responses is prob- ably brought about by the recognition of invariant pathogen-associated molecular patterns PAMP that are characteristic of whole classes of microbial organisms. Plants respond to pathogens by activating broad-spectrum innate immune responses that can be expressed locally at the site of pathogen invasion as well as systemically in the uninfected tissue.
Rotting of stored apples Malus domestica Borkh by Phlyctaena vagabunda Desm. Gloeosporium album Osterw Plate 1. An important characteristic of the fungus is that spores of P. There are conditions depend- ing on the fungus and the nature of vegetable tissue, in which infections, which take place in lenticels, can develop during storage to produce lesions. The available evidence Lattanzio et al. In vitro bioassays have shown that none of these naturally-occurring phenolics in concentrations like those encountered in fresh fruit exhibit inhibitory activity against spore germination or mycelial growth of P.
If pre-existing antifungal phenolics are not sufficient to stop the development of the infectious process, plant cells usually respond hypersensitive reaction by blocking or delaying the microbial invasion. Reactive oxygen species are often generated as warning signals within the cell or neighbouring cells, triggering off various reactions. These include the rapid increase of pre-existing antifungal phenols at the infection site, after an elicited increased activity of the key enzymes PAL and chalcone synthase of the biosynthetic pathway; this functions to slow or even halt the growth of the pathogen and to allow for the activation of secondary strategies that would restrict the pathogen more thoroughly.
PPO activity also increased in these tissues, to 2—3 times that in healthy tissues. Post-infection accumulation of pre-existing phenolics, especially phloridzin and chlorogenic acid which are good substrates of apple PPO, provides an adequate substrate to the increased PPO activity.
Thus, it cannot be excluded that after oxidative transformation phenolics are involved in induced resistance. The enzyme consumes oxygen and produces quinones or semiquinones, highly reactive compounds with potential toxic properties, and this makes the medium unfavourable to further development of pathogens Byrde et al. In vitro bioassays showed that, when a crude extract of apple PPO was added to a spore suspension of P.
These bioassays also showed a potential synergistic effect of phloridzin and chlorogenic acid Fig. Phloridzin alone oxidised slowly and formed the light yellowish reaction products. However, the simultaneous presence of chlorogenic acid in a model system increases the oxidation rate of phloridzin in the presence of PPO by decreasing the lag period of the enzymatic reaction. From these data, it appears that infection of apple tissue elicited an active glycosidase and PPO capable of converting phloridzin to phloretin, which was subsequently oxidised.
Simultaneously with hydrolysis to phloretin, phloridzin is oxi- dised via 3-hydroxyphloridzin to the corresponding o-quinone. The formed o-quinones are transient intermediates that may rapidly undergo oxidative condensation reactions Fig.
These transformation reactions of phloridzin in the presence of apple PPO indicate that oxidation products may be involved in the defence mechanism of apple against the fungus P. Chemical inhibitors also play an important role in the inhibition of oviposition on the host-plant, and, in turn, on insect larval growth and the survival of progeny.
Studies on the role of inhibitors in host plant selection indicate that many different chemicals may be expected to have an inhibitory effect on feeding by different insects. It is now generally accepted that plant phenolics play a role in protecting plants from insects Painter, ; Thorsteinson, ; Dethier, ; Chapman, ; Joerdens-Roettger, ; Ferguson et al. Some cotton flavonoids are feeding stimulants for the boll weevil, Anthonomus grandis Hedin et al. Nishida et al. Cowpea Vigna unguiculata L.
The major constraints to cowpea production are insect pests, plant diseases, plant parasitic weeds, drought and heat Murdock, ; Singh et al. Additional plant damage can also be caused by plant viruses that some aphid species transmit. There are two Aphis spp.
Homoptera: Aphididae reported as pests of cowpeas: Aphis craccivora Koch cowpea aphid , which is the main aphid infesting cowpeas throughout Africa and Asia, and Aphis fabae Scopoli black bean aphid , which has been reported as a minor pest in Africa and whose biology appears to be similar to that of A. Cowpea aphids primarily infest seedlings, but large populations also infest flowers and green pods of older plants Plate 1.
The resistant lines have a higher total flavonoid content than susceptible lines. This relationship was further confirmed when the flavonoid aglycone content of two near-isogenic lines of V. Fabae: quercetin is the most active whereas kaempferol has little effect on the reproduction rate.
Many flavonoids can act as feeding deterrents to phytophagous insects at relatively low concentrations. Therefore, the concentrations of flavonoids in plants are normally far higher than those needed for a deterrent effect on aphid feeding. Accession Daily larval deposition Kempferol chemotype: Vigna luteola Jacq. Bentham TVnu 7. Merrill var. Bentham TVnu Isorhamnetin chemotype: 0. There are kaempferol chemotypes, in which kaempferol is the only or the main aglycone detected, quercetin chemotypes, containing only quercetin glycosides, and isorhamnetin chemotypes.
From an ecological point of view, the most interesting chemotypes are some accessions, belonging to the same species, which make it possible to study, ceteris paribus, the role of endogenous flavonoids in plant resistance to aphids. Two chemotypes were found amongst Vigna marina accessions: V. When the resistance characteristics to aphids in different chemotypes of the same species were tested Table 1.
The cowpea seed beetle, Callosobruchus maculatus Fabricius Coleoptera: Bruchidae is a major pest of stored cowpeas, but actually infests the green pods while they are still in the field. Larvae hatch from eggs and penetrate the pod wall or the seed testa with their mouthparts. Therefore, the strong resistance of some cultivated or wild Vigna species to C. The resistance of TVu to bruchids was investigated by Gatehouse et al.
However, some researchers suggest that the trypsin inhibitor alone does not account for bruchid resistance in cowpea, thus indicating the need for further investigations. In addition, seed coat tannins are present at high levels in most plant seeds and grains, and are generally considered to be harmful to phytophagous insects.
Tannins may affect the growth of insects in three main ways: they have an astringent taste which affects palatability and decreases feed consumption; they combine with proteins to form complexes of reduced digestibility; and they act as enzyme inactivators Winkel-Shirley, In stored cowpea, seed coat proanthocyanidins contribute to resistance against cowpea weevil C.
On the contrary, the seed coat tannin content was found to be 13 times higher in undamaged Vita 7 seeds than in IT 84E- infested seeds. These results support the hypothesis that, if bruchids infest cowpea when the grain is stored after harvest, seed coat tannins are effectively involved in the biochemical defence mechanisms, which can deter, poison or starve the bruchid larvae that feed on cowpea seeds.
This chemical response to changing environments has led to the enormous structural variation in the major groups of phenolic compounds, which are evident in plants today. More detailed knowledge of these effects should enable prediction and selection of growth conditions in order to achieve a desirable content of these secondary metabolites. Manipulation of environmental factors should — at least to some degree — represent an alternative to genetic engineering for achieving special effects on the level of plant components.
Furthermore, understanding of the regulatory and biochemical mechanisms that control the types and amounts of phenolic compounds synthesised under different conditions continues to be a high priority for research, with a view to possible engineering of crop plants to overproduce antioxidant phenolics.
Broadly speaking, plant growth and productivity are greatly affected by environmen- tal stresses. Both abiotic and biotic stresses divert substantial amounts of substrates from primary metabolism into secondary defensive product formation and this could lead to constraints on growth.
Plants have limited resources to support their physiological pro- cesses, so that all requirements cannot be met simultaneously, and trade-offs occur between growth and defence Coley et al. Therefore, a principal feature of plant metabolism is the flexibility to accommodate devel- opmental changes and respond to the environment.
The cellular and molecular responses of plants to environmental stress include mechanisms by which plants perceive environmental signals and transmit the signals to cellular machinery to activate adaptive responses, and this is of fundamental importance to biology. Knowledge about stress signal transduction is also vital for the continued development of strategies to improve stress tolerance in crops Xiong et al. In addition, plant responses to both biotic and abiotic stresses require the reallocation of resources, therefore these responses are considered to be costly for plants because of the energy consumed in the biosynthesis of defensive phenolics and the ecological consequences of their accumulation.
Costs can be described as resource-based trade-offs between resistance and fitness, as ecological costs, or as allocation costs Heil et al. In order to quantify these costs in plants, researchers have attempted to link a measure of plant success usually, growth rate with levels of defensive compounds. Zangerl et al. Pavia et al. Data showed that there was a significant negative relationship between phlorotan- nins and growth. Resource-based alloca- tion theory predicts a trade-off mechanism between plant reproduction, growth and defence functions that regulates carbon fluxes between primary and secondary metabolism, and that is specifically required for protective adaptation to environmental stresses Coley et al.
In many plants, free proline also accumulates as a common physiological response to a wide range of biotic and abiotic stresses. Furthermore, proline accumulation is consid- ered to be one of the stress signal influencing adaptive multiple responses that are part of the adaptation process.
Transgenic approaches have confirmed the beneficial effect of proline overproduction during stress. Accumulation of proline could be due to de novo synthesis, to decreased degradation, or to both of these.
Is there a link between increased phenolic levels and increased proline levels in plant tissues under stress? In this connection, it must be stressed that the oxidative pentose phosphate pathway OPPP is the source of reducing equivalents NADPH for phenylpropanoid biosynthesis, and that this pathway also provides the ery- throse 4-phosphate that, along with phosphoenolpyruvate formed from glycolysis, serves as a precursor for phenylalanine biosynthesis via the shikimic acid pathway Fahrendorf et al.
A fixed amount of resources is usually assumed to be divided among fixed maintenance cost, growth and reproduction. Growth rates are correlated with the ecological conditions in which each species is living in nature; slow growth is adaptive for dealing with environmental stresses.
Growing plants, therefore, continuously face a dilemma regarding the partitioning of their available carbon resources. If priority is given to the plant growth processes, the availability of carbon resources and other nutrients may become limiting for plant defence-related processes, and vice versa.
So far, four main plant defence hypotheses have been put for- ward to explain patterns and variations in the concentration of carbon-based secondary compounds in plant tissues, according to availability of resources. These theories hinge on the presence of resistance costs, because, in the absence of costs, selection is expected to favour the best-defended genotypes.
This hypothesis also suggests that carbon-based secondary metabolites tend to accumulate when growth is limited by low levels of mineral nutrients. The optimal defence theory ODT; McKey, has served as the main framework for investigation of genotypic expression of plant defence, with the emphasis on the allocation cost of defence.
This theory addresses how the defensive needs of a plant contribute to the evolution of secondary metabolites, with defence costs paid to maximise plant fitness. In essence, this hypothesis states that any defensive pattern is possible if it is adaptive. PCM states that protein and phenol synthesis compete for the common, limiting resource phenylalanine, so that protein and phenolic allocation are inversely correlated. Phenol allocation can be predicted from the effects of development, inherent growth rate and environment on leaf functions that create competing demands for proteins or pheno- lics.
Allocation to differentiation includes the cost of enzymes, transport and storage structures involved in defence. Growth and secondary metabolism can compete for available photosynthates and so there is a trade-off for carbon allocation.
GDBH states that there is a physiological trade-off between growth and secondary metabolism imposed by developmental constraints in growing cells, and competition between primary and secondary metabolic pathways in mature cells. This hypothesis also predicts that any factor that slows growth more than it slows photosyn- thesis can increase the internal resources available for allocation to differentiation.
For instance, growth is slowed by the limitation of nutrients, whereas photosynthesis is less sensitive to it. Consequently, carbohydrates accumulate beyond growth demands, and may thus be converted to secondary metabolites. In nature, cellular functions are propagated by cascades of molecules, which interact with one another. Generally speaking, one reaction depends on a previous step.
These data are consistent with the scheme proposed in Fig. Plant tissues are forced to accumulate free proline under stress conditions. Rather, it is rapidly recycled back to regenerate phenylalanine, thereby providing an effective means of maintaining active phenylpropanoid metabolism with no additional nitrogen requirement. Finally, in good agreement with the scheme proposed in Fig. References Abdel-Farid, I. Plant Science, , — Adams-Phillip, L. Plant Physiology, , — Alibert, G.
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