Towards understanding factors that govern fertilizer response in casave: lessons from East Africa

Information on fertilizer response in cassava in Africa is scarce. We conducted a series of on-farm and on-station trials in two consecutive years to quantify yield responses of cassava to mineral fertilizer in Kenya and Uganda and to evaluate factors governing the responses. Average unfertilized yields ranged from 4.2 to 25.7 t ha-1 between sites and years. Mineral fertilizer use increased yields significantly, but response to fertilizer was highly variable (-0.2 to 15.3 t ha-1). Average yield response per kg applied nutrient was 37, 168 and 45 and 106, 482 and 128 kg fresh yield per kg of applied N, P and K, respectively in 2004 and 2005. Fertilizer response was governed by soil fertility, rainfall and weed management, but was not influenced by variety, pest and disease pressure and harvest age. Relative N and K yields were positively correlated to SOC and exchangeable K, while response to fertilizer decreased on more fertile soils. Still, fertilizer response varied widely on low fertility soils (e.g. on soils wit

A comparative study of the physicochemical properties of starches from root, tuber and cereal crops

Some properties of starches from cassava, potato and sweet potato were compared with cereal starches from maize, wheat, millet and sorghum. The aim was to determine the properties of tuber and root crop starches and compare them with cereal starches in addition to unravelling the potential of commonly grown sorghum and millet climate resilient crops as cheap and sustainable sources of starch. Significant variations were observed for amylose content and solution properties of starches, where blue values for amylose ranged from 0.355 in potato to 0.476 in cassava, but were averagely low in cereal starches. Amylose leaching increased with temperature with the highest value (0.432) in cassava at 80°C compared with cereal starches (average 0.361). Starch amylosis increased with time of hydrolysis and was highest (>16%) for millet and sorghum and least for potato (<8.5% average). Average swelling power at 80°C was high for cassava (8.58 g/g) and potato (8.44 g/g) compared with sweet potato (6.88 g/g) and low among cereal starches (5.17 g/g). Similarly, starch solubility was low in potato (0.77 g/g) and sweet potato (0.577 g/g) compared with cassava (1.23 g/g). The paste clarity was also high for cassava (48.32%) and potato (42.16%) and least for sweet potato derived starches (23.22%) and all the cereal starches (14.97%). These properties demonstrate the untapped potential of cassava and tuber based starches for use in food and non-food applications previously dominated by cereal starches.Key words: Tuber starch, root crop starches, cereal starches, amylose, amylosis

Production of friable embryogenic callus and regeneration of Ugandan farmer-preferred cassava genotypes

Generation of embryogenic callus is a key step in genetic engineering of many crop species, including cassava. Protocols for generation of friable embryogenic callus (FEC) have been lacking for Ugandan cassava genotypes, thereby delaying their genetic engineering for agronomic and other desirable traits. The objective of this study was to determine conditions suitable for production and regeneration of FEC in the Ugandan cassava genotypes; Aladu, Bukalasa and Ebwanateraka, and control cultivar 60444. Immature leaf lobe explants were established on Murashige and Skoog (MS) based media for initiation of organized embryogenic callus (OES). To produce FEC, resulting OES were established on Gresshoff and Doy based callus induction media with varying levels of sucrose, maltose, tyrosine, tryptophan, naphthalene acetic acid (NAA) under light and dark conditions. Subsequently, FEC was subcultured to MS-based embryo maturation and embryo regeneration media. All genotypes produced OES. All genotypes produced FEC except Bukalasa. The amino acid tyrosine favoured production of FEC in Aladu and Ebwanatereka, but not in 60444, while 20 g/L of sucrose trigged production of FEC in Aladu and 60444, but 40 g/L of sucrose was superior for Ebwanatereka. Media supplemented with 1 ml/L naphthalene acetic acid NAA facilitated embryo regeneration in Ebwanatereka and 60444, while Aladu responded better to 5 ml/L NAA. Light, tyrosine and sucrose were essential for FEC production in Uganda cultivars while NAA was required for regeneration of somatic embryos. Ability to produce FEC in these genotypes lays a foundation for their improvement through genetic transformation for the desired and agronomic traits.Key words: Cassava (Manihot esculenta Crantz), somatic embryogenesis, amino acids, carbon sources

Morphotypes, distribution and uses of false banana in Uganda

Enset ( Ensete ventricosum ) is commonly known as the \u201cfalse banana\u201d because of its close resemblance to the domesticated banana ( Musa spp.) plant. It is Ethiopia\u2019s most important traditional staple crop in the densely populated south and south-western parts of the country; where it is grown and exploited for its starch to make various food and industrial products. In Uganda, little is known about enset regarding its ethno-botany and distribution, yet it occurs in the country. The objective of this study was to map out the distribution and document the ethno-botany and uses of enset in Uganda. A survey was carried out throughout the country to identify its natural habitats in different regions. Local people were interviewed on the uses of enset, etymology, and identification of different morphotypes. Morphological descriptors and sex of enset accessions were used in classification or identification of morphotypes. Enset was generally widely, but sparsely distributed in the different regions in Uganda; growing at elevations ranging between 988 (Moyo district) to 2,150 (Kapchorwa district) metres above sea level (masl) and in a variety of habitats. Out of the 80 districts of Uganda (as of 2009), enset was reported and observed in 30 districts. Thirteen local names of enset and their meanings were documented; but it was widely referred to as Kitembe. Different plant parts were used for medicinal purposes; while the leaves were used in local beer brewing. There were five enset morphotypes distinguished by morphological traits, such as plant height (short vs tall), mid-rib colour (light green vs pink), pseudostem background appearance (light green vs brown), leaf margin colour, male bud colour, and leaf length-breadth ratio. A detailed molecular level genetic diversity assessment is recommended for further validation of the morphotypes.Le Bananier d\u2019Abyssinie ( Ensete ventricosum ) est commun\ue9ment appel\ue9 comme \uab\ua0fausse banane\ua0\ubb \ue0 cause de sa tr\ue8s grande ressemblance au plant de banane ( Musa spp) domestiqu\ue9. C\u2019est un aliment de base traditionnel tr\ue8s important dans les r\ue9gions de grande densit\ue9 de population du Sud et Sud-Ouest d\u2019Ethiopie\ua0; o\uf9 il est cultiv\ue9 et exploit\ue9 pour son amidon utilis\ue9 pour divers produits alimentaires et industriels. En Ouganda, tr\ue8s peu d\u2019informations relatives \ue0 son ethnobotanique et \ue0 sa distribution sont disponibles sur Bananier d\u2019Abyssinie, pourtant il est pr\ue9sent dans le pays. L\u2019objective de cette \ue9tude \ue9tait de cartographier la distribution et de documenter l\u2019ethnobotanique et les usages du bananier d\u2019Abyssinie en Ouganda. Une enqu\ueate a \ue9t\ue9 conduite \ue0 travers le pays pour identifier ses habitats naturels dans diff\ue9rentes r\ue9gions. Les populations locales ont \ue9t\ue9 interview\ue9es sur les usages du bananier d\u2019Abyssinie, l\u2019\ue9tymologie, et l\u2019identification des diff\ue9rents morphotypes. Les descripteurs morphologiques et le sexe des accessions du bananier d\u2019Abyssinie ont \ue9t\ue9 utilis\ue9s dans la classification ou l\u2019identification des morphotypes. Le bananier d\u2019Abyssinie a \ue9t\ue9 largement, but sporadiquement distribu\ue9 dans les diff\ue9rentes r\ue9gions en Ouganda\ua0; les altitudes de cultures varient entre 988 (district de Moyo) \ue0 2150 m\ue8tres (district de Kapchorwa) au-dessus du niveau de la mer (masl) et dans une vari\ue9t\ue9 d\u2019habitats. Sur les 80 districts (comme en 2009), le bananier d\u2019Abyssinie a \ue9t\ue9 rapport\ue9 et observ\ue9 dans 30 districts. Les treize noms locaux du bananier d\u2019Abyssinie et leurs significations ont \ue9t\ue9 document\ue9s\ua0; mais il a \ue9t\ue9 largement r\ue9f\ue9r\ue9 au Kitembe. Les diff\ue9rentes parties du plant ont \ue9t\ue9 utilis\ue9es \ue0 des fins m\ue9dicinales, alors que les feuilles ont \ue9t\ue9 utilis\ue9es dans la pr\ue9paration de la bi\ue8re locale. Il y avait cinq morphotypes du bananier d\u2019Abyssinie distingu\ue9s par des traits morphologiques, tels que la hauteur du plant (courte vs haute), couleur de mid-nervure (vert-clair vs rose), fond d\u2019apparence de pseudo tige (vert-clair vs marron), couleur de la marge de la feuille, m\ue2le mais la couleur, et un ratio longueur/largeur de la feuille. Une \ue9valuation d\ue9taill\ue9e de la diversit\ue9 g\ue9n\ue9tique mol\ue9culaire est recommand\ue9e pour davantage validation des morphotypes

Influence of spectral properties on cassava leaf development and metabolism

Cassava’s importance as a food security crop in Sub Saharan Africa will be enhanced by its special traits such as tolerance to drought and high yields under hydrothermal stress. Some of the special traits which include the light reflective and absorptive properties of the leaves that depend on the surface characteristics of the leaves, are variety dependent and may influence the plants’ reaction to light; hence, its photosynthetic capacity. We investigated the differences in the leaf spectral properties in different cassava varieties and related them to leaf biochemical properties using 20 cassava varieties established in a randomized complete block design in Kasese, western Uganda. Time dependent changes in leaf spectral characteristics were studied using Digimizer software and related to changes in sugar and pigment properties. Changes in the amount of reflected light were observed for the three main wavelengths used by plants (blue, green and red) with the blue being the most preferred. Total soluble free sugars exhibited a diurnal pattern from lower values (0.07 mg/g) after the dark period to higher values (0.313 mg/g) as the day progressed and was different from those of translocatable sugars such as sucrose. Chlorophyll a exhibited a curved pattern in all varieties increasing with increase in light intensity from 09:00 h (0.18 ug/g), peaking at 15:00 h (0.22 ug/g) and dropping down in concentration by 18:00 h (0.16 ug/g). Significant differences were observed in cassava varieties for the concentration of chlorophylls and carotenes. The results were obtained at a time of optimal growth conditions (four months after planting) and were used to classify these varieties into three broad groups showing that studies on spectral properties of leaves can still give a lot of insights in selection for stress tolerance under less optimal stress. The significant changes observed in the phenotype especially the foliar portion of the plant with the stay green and early recovering mechanisms of tolerance identified also tarried well with observed spectral differences. The results show that studies on plant spectral properties can be important in making inferences on the plants physiological and growth status.Keywords: Spectral properties, tolerance mechanisms, physiology, reflectance

Effect of self-pollination with heat-treated pollen on parthenocarpy and homozygosity in cassava

Cassava\u2019s ( Manihot esculenta Crantz) high heterozygosity complicates its genetic improvement via selective breeding. Double haploid (DH) technology can be used to improve the crop\u2019s heterozygosity, thereby improving the capacity for genetic improvement. The objective of this study was to evaluate the effect of self-pollination using heated pollen on pollen tube penetration, fruit set, seed and haploid embryo development in cassava genotypes for the production of haploid cassava. Pollen from two cassava genotypes, NASE3 and NASE14, was heated at 40, 50 and 60 oC for 0.5, 1.0 and 2.0 hr each. The heated pollen was used in six rounds of self-pollinations. Pollen tube penetration was monitored by fluorescent microscopy, followed by early embryo rescue and ovule culture. Ploidy and zygosity were assessed using flow cytometry and single-nucleotide polymorphism analysis, respectively. Pollen germinated on the stigma, grew within the style through the nucellar beak, but did not reach the embryo sac, thus achieving no fertilisation in all the 5756 self-pollinated flowers. There was a reduction in pollen germination (in vitro and in vivo), pollen tube penetration and fruit set with increasing temperature. Heat-treated pollen stimulated division of the egg cell and induced development of parthenocarpic fruits. Up to 6 embryoids per ovule were observed and all regenerated plantlets were diploid, with up to 93.0% increased homozygosity. For the first time, plant regeneration from ovules, pollinated with fresh pollen at 14 days after pollination, was achieved indicating improved speed in plant regeneration. The data generated are important for the development of protocols for cassava DH plant production.La forte h\ue9t\ue9rozygotie du manioc ( Manihot esculenta Crantz) complique son am\ue9lioration g\ue9n\ue9tique par s\ue9lection s\ue9lective. La technologie d\u2018 haplo\uefde double (DH) peut \ueatre utilis\ue9e pour am\ue9liorer l\u2019h\ue9t\ue9rozygotie de la culture, am\ue9liorant ainsi la capacit\ue9 d\u2019am\ue9lioration g\ue9n\ue9tique. L\u2019objectif de cette \ue9tude \ue9tait d\u2019\ue9valuer l\u2019effet de l\u2019auto-pollinisation \ue0 l\u2019aide de pollen chauff\ue9 sur la p\ue9n\ue9tration du tube pollinique, la nouaison, le d\ue9veloppement des graines et des embryons haplo\uefdes dans les g\ue9notypes de manioc pour la production de manioc haplo\uefde. Le pollen de deux g\ue9notypes de manioc, NASE3 et NASE14, a \ue9t\ue9 chauff\ue9 \ue0 40, 50 et 60 oC pendant 0,5, 1,0 et 2,0 heure (s) chacun. Le pollen chauff\ue9 a \ue9t\ue9 utilis\ue9 dans six cycles d\u2019auto-pollinisation. La p\ue9n\ue9tration du tube pollinique a \ue9t\ue9 surveill\ue9e par microscopie fluorescente, suivie d\u2019un sauvetage pr\ue9coce des embryons et d\u2019une culture d\u2019ovules. La plo\uefdie et la zygosit\ue9 ont \ue9t\ue9 \ue9valu\ue9es \ue0 l\u2019aide de la cytom\ue9trie en flux et de l\u2019analyse du polymorphisme mononucl\ue9otidique, respectivement. Le pollen a germ\ue9 sur le stigmate, s\u2019est d\ue9velopp\ue9 dans le style \ue0 travers le bec nucellaire, mais n\u2019a pas atteint le sac embryonnaire, n\u2019obtenant ainsi aucune f\ue9condation dans toutes les 5756 fleurs autogames. Il y avait une r\ue9duction de la germination du pollen (in vitro et in vivo), de la p\ue9n\ue9tration du tube pollinique et de la nouaison avec l\u2019augmentation de la temp\ue9rature. Le pollen trait\ue9 thermiquement a stimul\ue9 la division de l\u2019ovule et induit le d\ue9veloppement de fruits parth\ue9nocarpiques. Les 6 embryo\uefdes par ovule ont \ue9t\ue9 observ\ue9s et toutes les plantules r\ue9g\ue9n\ue9r\ue9es \ue9taient diplo\uefdes, avec 93,0% d\u2018augmentation d\u2019homozygotie. Pour la premi\ue8re fois, la r\ue9g\ue9n\ue9ration des plantes \ue0 partir des ovules, pollinis\ue9es avec du pollen frais 14 jours apr\ue8s la pollinisation, a \ue9t\ue9 r\ue9alis\ue9e, indiquant une vitesse am\ue9lior\ue9e de r\ue9g\ue9n\ue9ration des plantes. Les donn\ue9es g\ue9n\ue9r\ue9es sont importantes pour l\u2019\ue9laboration de protocoles de production de plantes de manioc de DH

Diurnal oscillation of SBE expression in sorghum endosperm

Spatial and temporal expression patterns of the sorghum SBEI, SBEIIA and SBEIIB genes, encoding, respectively, starch branching enzyme (SBE) I, IIA and IIB, in the developing endosperm of sorghum (Sorghum bicolor) were studied. Full-length genomic and cDNA clones for sorghum was cloned and the SBEIIA cDNA was used together with gene-specific probes for sorghum SBEIIB and SBEI. In contrast to sorghum SBEIIB, which was expressed primarily in endosperm and embryo, SBEIIA was expressed also in vegetative tissues. All three genes shared a similar temporal expression profile during endosperm development, with a maximum activity at 15-24 days after pollination. This is different from barley and maize where SBEI gene activity showed a significantly later onset compared to that of SBEIIA and SBEIIB. Expression of the three SBE genes in the sorghum endosperm exhibited a diurnal rhythm during a 24-h cycle

Fruit set and plant regeneration in cassava following interspecific pollination with castor bean

The increasing demand for cassava ( Manihot esculenta Crantz) for food and non-food uses in the tropics necessitates that its breeding for increased root productivity be made faster. The characteristic long breeding cycle and heterozygous nature of this crop, pose a major obstacle to its rapid genetic improvement. This study aimed at inter-pollinating cassava with castor bean ( Ricinus communis ), with a purpose of inducing and regenerating cassava doubled haploids (DHs). A total of 3,349 flowers from twelve elite cassava varieties were inter-pollinated with caster bean. A total of 803 fruits were harvested for early embryo rescue and/or ovule culture. Of these, three were dissected to obtain seven unique embryos, while 800 were dissected to obtain 1312 young ovules, all of which were cultured in vitro. Overall, 82 (6.25%) of the cultured ovules formed callus that originated from the embryosac region, which is haploid. Four out of seven rescued embryos (57.1%) regenerated into plantlets. Ploidy analyses of 24 samples using flow cytometry revealed that 23 of the analysed samples were diploid. However, one callus sample was anueploid. Only one sample had an exceptionally high level of homozygosity ( 84.2%). These findings lay a foundation for future research aimed at induction of haploids in cassava.La demande croissante de manioc (\ua0 Manihot esculenta \ua0Crantz\ua0) \ue0 usage alimentaire et non alimentaire dans les tropiques\ua0n\ue9cessite que sa reproduction pour une productivit\ue9 accrue des racines soit faite plus rapidement.\ua0Le long cycle de reproduction et le caract\ue8re h\ue9t\ue9rozygote de cette plante constituent un obstacle majeur dans la rapidit\ue9 de son am\ue9lioration g\ue9n\ue9tique.\ua0Cette \ue9tude visait \ue0 inter-polliniser le manioc avec le haricot (\ua0 Ricinus communis \ua0), dans le but d\u2019induire et de r\ue9g\ue9n\ue9rer le manioc d\u2019haplo\uefdes doubl\ue9 (HD).\ua0Un total de 3 349 fleurs de douze \ue9lites vari\ue9t\ue9s de manioc ont \ue9t\ue9 inter-pollinis\ue9es avec le haricot.\ua0Un total de 803 fruits ont \ue9t\ue9 r\ue9colt\ue9s pour les embryons pr\ue9matur\ue9s qui etaient sauves et\ua0/ ou la\ua0culture d\u2019ovules\ua0.\ua0Parmi ceux-ci,\ua0trois ont \ue9t\ue9 diss\ue9qu\ue9s pour obtenir sept embryons uniques\ua0,\ua0tandis que 800 ont \ue9t\ue9 diss\ue9qu\ue9s pour obtenir 1312 jeunes ovules, qui ont tous \ue9t\ue9 cultiv\ue9s\ua0in vitro\ua0.\ua0Un total de 82 (6,25%) des ovules en culture ont form\ue9 des cals provenant de la\ua0r\ue9gion\ua0embryonnaire\ua0, qui est haplo\uefde. Quatre parmi sept embryons sauv\ue9s (57,1%) se sont r\ue9g\ue9n\ue9r\ue9s en plantules.\ua0Les analyses de plo\uefdie de 24 \ue9chantillons par cytom\ue9trie en flux ont r\ue9v\ue9l\ue9 que 23 des \ue9chantillons analys\ue9s \ue9taient diplo\uefdes.Cependant, un \ue9chantillon de cals \ue9tait anueplo\uefde.\ua0Un seul \ue9chantillon pr\ue9sentait un niveau d\u2019homozygotie exceptionnellement \ue9lev\ue9\ua0(84,2\ua0%).\ua0Ces r\ue9sultats sont les bases des recherches dans le futur sur la cause des haplo\uefdes dans le manioc

In vitro embryo rescue and plant regeneration following self-pollination with irradiated pollen in cassava (Manihot esculenta Crantz)

Cassava is a highly heterozygous species; hence, current methods used in classical cassava breedingcannot match the urgent need to high yielding varieties. Recently, progress was made through androgenesis and gynogenesis as pathways for raising doubled cassava haploid lines to overcome problems associated with cassava’s inherent reproductive biology, but these efforts were limited (nocandidate cassava plantlets were regenerated). For the first time, this study shows that pollen irradiation coupled with self-pollination and embryo rescue regenerated 62 candidate cassava plantlets. Plants of an elite cassava variety, Nase14, served as a mother plant and as the pollen donor for the irradiation. Irradiation dosages of 50 to 250 Gray studied across five pollination events and 300 or 500 Gray in one pollination event caused a reduction in pollen germination up to 67.0%. By 15 days after pollination (DAP) with irradiated pollen, up to 89.7% of the pollinated flowers had aborted. By embryo rescue time (42 DAP), significant differences were observed in number of fruits, seeds and embryos generated, with the non-irradiated pollen treatments having significantly higher numbers. Sixteen (16) heterozygous SSR markers in the parent and ploidy analysis showed that none of the regenerated plants was haploid or homozygous. However, the plantlets resulting from pollination with non-irradiated pollen had 56.2% homozygous loci, while progeny derived from irradiated treatments had frequencies of homozygous loci between 28.1 and 55.0%. This is the first time to use irradiated pollen in cassava as a pathway to generate candidate plantlets as an initial step in double haploid production.Key words: Cassava, doubled haploids, embryo rescue, plant regeneration, pollen germination, pollenirradiation

CASSAVA BROWN STREAK DISEASE EFFECTS ON LEAF METABOLITES AND PIGMENT ACCUMULATION

Cassava Brown Streak Disease (CBSD) is a threat to productivity and product quality in East Africa. The objective of this study was to understand the effect of CBSD on the primary photosynthetic apparatus of cassava ( Manihot esculenta Crantz). Three cassava varieties with varying levels of reaction to infection by CBSD were chosen and subjected to field disease pressure. Disease progression and the resulting effects on leaf morphology, metabolite and pigment accumulation were assessed over a period of five months, beginning from 3 months after planting. Slight increments in leaf dry matter were observed up to 4 MAP, and there after a drop was registered. A significant (P<0.05) reduction in photosynthetic pigments occured with a fall in the Chla:Chlb and Chla:Carotenoid ratio, indicative of specific reductions in chlorophyll a (-80%) compared to Chlorophyll b (-41 to -62%) and Carotenoid (-11 to -18%). Total reducing sugar and starch content also dropped significantly (-30 and -60%, respectively), much as NASE 14 maintained a relatively higher amount of carbohydrates. Leaf protein levels were significantly reduced at a rate of 0.07 ug g-1 leaf per month in diseased treatments. Significant reductions in primary metabolites show altered leaf photosynthetic and growth metabolism, resulting into a compromised plant system that cannot perform optimally.La Maladie Stri\ue9e Brune du Manioc (CBSD) constitue une menace pour la productivit\ue9 et la qualit\ue9 des produits du manioc en Afrique de l\u2019Est. L\u2019objectif de cette \ue9tude \ue9tait de comprendre l\u2019effet de CBSD sur l\u2019appareil photosynth\ue9tique primaire du manioc ( Manihot esculenta Crantz). Trois vari\ue9t\ue9s de manioc poss\ue9dant differents niveaux de r\ue9sistance \ue0 CBSD ont \ue9t\ue9 soumises \ue0 la pression dans les conditions de plein champs. La progression de la maladie et son effet sur la morphologie foliaire, les metabolites et accumulations de pigments ont \ue9t\ue9 \ue9valu\ue9s sur une p\ue9riode de cinq mois, \ue0 compter du troisi\ue8me mois apr\ue8s semis. Il a \ue9t\ue9 observ\ue9e une l\ue9g\ue8re augmentation de mati\ue8re s\ue8che foliaire jusqu\u2019\ue0 4 MAP, suivie d\u2019une chute. Une r\ue9duction significative (P<0,05) des pigments photosynth\ue9tiques a \ue9t\ue9 observ\ue9e avec une chute dans le Chla:Chlb et Chla:ratio de Carotenoide, indiquant des r\ue9ductions sp\ue9cifique sen chlorophyllea (-80%) en comparaison avec la Chlorophylleb (-41 \ue0 -62%) et les Carotenoides (-11 \ue0 -18%). La r\ue9duction du sucre et d\u2019amidona connu aussi une diminution significative (-30 et -60%, respectivement), de m\ueame NASE 14 a maintenuun equantit\ue9 relativement\ue9 lev\ue9e d\u2019hydrates de carbone. Les niveaux de proteines foliaires \ue9taient significativement r\ue9duits \ue0 0,07 ug g-1 de feuille malade par mois. Des r\ue9ductions significatives des metabolites primaires sont \ue0 l\u2019origne de l\u2019alt\ue9ration du m\ue9tabolisme photosynth\ue9tique et de croissance, resultant \ue0 cycle v\ue9g\ue9tatif compromis qui ne peut permettre un d\ue9veloppement optimal de la plante