Tolerance of Different Host-Plants to the Cassava Mealybug Phenacoccus Manihoti Mati Le-Ferrero (Homoptera: Pseudococcidae)
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Tolerance of different host-plants to the cassava mealybug Phenacoccus manihoti Mati le-Ferrero (Homoptera: Pseudococcidae) (Keywords: Phenacoccus manihoti, cassava, insect-plant relationship, antibiosis, tolerance, varietal resistance, Congo) B. LE RU and M. TERTULlANO Laboratoire d'Entomologie Agricole, ORSTOM, BP 181, Brazzaville, R. du Congo Abstract. The tolerance of seven varieties of cassava (Manihot Aphididae (Dixon, 1971 a,b; Barlow et al., 1977; Barlow esculenta Crantz), Poinsettia (Euphorbia ptncbetrime Wild.), and Messmer, 1982; Hawkins et al., 1985, 1986). Their Talinum (Talinum triangulare Jacq.) and Faux-caoutchouc (a quantification provides a useful tool to evaluate the degree hybrid of M. esculenta and M. glaziovii Mull. Arg.) to the cassava of plant tolerance (Jimenez et al., 1989; Scott et aI., 1991; mealybug Phenacoccus manihoti Matt. Ferr., was measured in the laboratory, by quantifying morphological and physiological Webster et al., 1991). changes in the growth of plants infested with P. menihotl. A 2 When evaluating components of host-plant resistance month infestation by 100 mealybugs decreased the total leaf area to P. manihoti, different degrees of resistance in terms of and the leaf area ratio but did not modify total dry weight, F, antixenosis and antibiosis have been demonstrated (Le relative growth rate R or net assimilation rate E of plants studied. Ru et al., 1991 a). The aim of the present study is to The decrease in leaf area was extremely variable and depended on plant species and variety. The effect of infestation on dry determine the cassava tolerance to P. manihoti by quan weight of different plant organs was not homogeneous, suggest tifying morphological and physiological changes in their ing that translocation of nutrients made during photosynthesis growth caused by pest infestation. was modified. In cassava the decrease in leaf area, without significant change in leaf dry weight, appeared to be due to physiological changes in plants similar to those induced by water stress. The degree of tolerance was not related to the plant 2. Materials and methods species (cassava varieties Ganfo, Incoza and Zanaga were as tolerant as Talinum and Faux-caoutchouc). The 10 host-plants used in this study were selected according to the results obtained in a previous experi ment done to assess the antixenotic and antibiotic rela tionships between P. manihoti and its host-plants (Le RO 1. Introduction et al., 1991 a). The seven varieties of cassava (M. esculenta) have a varying degree of resistance in terms of antixenosis Tolerance is a component of plant resistance and re and antibiosis (high resistance: Incoza, Moudouma, and lates to its capacity to withstand insect attacks which Zanaga; medium resistance: M'Pembe, 30 M7; low resist would otherwise affect the growth of susceptible plants ance: 3 M8, Ganfo). (Horber, 1980). Unlike antixenosis and antibiosis, it does We also measured the tolerance of Faux-caoutchouc (a not exert a strong selective pressure on insect pest popu hybrid of M. esculenta and M. glaziovii Mull. Arg.), Tall lations; thus it can be maintained in a population over a num (Talinum triangulare Jaq., Portulacacae) and Poin long period (Auclair, 1989; Jimenez et al., 1988). settia (Euphorbia pulcherrima Wild., Euphorbiaceae); Cassava (Manihot esculenta Crantz; Euphorbiaceae) is their biological characteristics are as follows: a perennial shrub plant, relatively tolerant to diseases and pests because, unlike most other cultivated plants, it does 1. Faux-caoutchouc hosts large populations of P. not have critical phases during its development which can manihoti throughout the year, unlike cassava on possibly affect the yield. However, yield losses are high if which they are present for only 3-5 months the lifespan of leaves and the rate of photosynthesis are (Iheagwam, 1981). decreased or if stems are damaged severely (Cock, 1979). 2. Talinum is a weedy plant found in cassava fields, and A field experiment done in Nigeria has shown that the can sometimes host large populations of cassava infestation of cassava by the mealybug Phenacoccus mealybugs (Neuenschwander et et., 1986). manihoti Matile Ferrero (Homoptera; Pseudococcidae) 3. Poinsettia is sometimes used in the laboratory as results in a yield loss ranging between 52% and 58% in a substitute for cassava to mass-rear mealybugs 12-month-old plants. This loss might be due to a decrease (A. Panis and J. Boussienguet, personal communica in growth rate and leaf area; the latter might strongly tion), although it is never infested with this pest influence the total production of dry matter and its redis under natural conditions in Congo. tribution to roots (Schulthess et al., 1991). These modifications in growth due to the infestation of Plants were obtained from 30 cm cuttings planted vertically P. manihoti are similar to those induced by other sucking with two-thirds of their length in soil held in plastic pots insects such as Cicadellidae (Andrzejewska, 1967) and (30 x 22 cm). Pots were left in the shade for 2 weeks to 0967-0874/93 $10'00 © 1993 Taylor and Francis Lld stimulate the rooting process. Thereafter they were placed where W o and W 1 are the total dry weights at the begin in full light untit the 9-1 O-Ieaf stage (about 70 cm in height) ning (To) and end (Td of the time period; R indicates the which took 9-10 weeks. Pots (30 of each host-plant) were variation in plant weight per unit of time. then left for a further 2 months in a growth chamber under The rate of net assimilation Ë is: controlled conditions with an average temperature of 25°C (minima-maxima 21-32°C) and an average relative humidityof 70% (ranging between 60% and 80%); photo where A o and A 1 are the leaf areas at times To and T 1 period was 12 hours L:D. Under these experimental con respectively; Ë is an approximate measure of the net ditions we obtained after a 4-month period, plants with photosynthetic rate (Causton and Venus, 1981). In a con approximately 20 leaves and about 90-100 cm in height. stant environment, Ë can be considered as an index of the Under natural conditions in Congo, attacks by mealybugs plant's productive efficiency (Williams, 1946). occur at the beginning of the dry season on plants 3-8 The leaf area ratio Fis: months otd derived from the propagation of cassava cut tings, mostiy performed between November and April. Pots were watered twice a week during ail the experiment. This is a morphological index of the plant, and assumes The strain of P. manihoti (species reproducing by thely that leaves are the sole assimilatory organs (Causton and tocous parthenogenesis) used for the experiment in the Venus, 1981). laboratory was collected on the M'Pembe variety in the Mean values were analysed statistically using at-test at vegetable crop area of Brazzaville in October 1988. In the 5% level. order to Hmit the influence of parental trophic feeding mealybugs were reared for four generations (6 months) on 3. Results and Discussion each of the host-plants used in this experiment. At time 0 (To), 15 plants per host-plant, of identical size, Infestation of P. manihoti decreased the total leaf area of without symptoms of African cassava mosaic virus, were ail host-plants studied; however, this decrease was signi chosen amongst the 30 cuttings prepared initially. These ficant only for the Faux-Caoutchouc and for varieties of were divided randomly into two groups: one group of five cassava Moudouma, Ganfo Zanaga, 30 M7 and M'pembe. plants which was harvested 2 months later at time T1 • Infestation also reduced significantly the average number Each pair consisted of a control plant (non-infested) and a of leaves in the varieties of cassava 3 M8 and 30 M7, as weil treated plant (infested with 100 neonate mealybug L1 as the average number of internodes in the variety 3 M8 larvae, less than a day old). These conditions of infestation and Talinum, and the stem length between internodes in (100 mealybugs per plant for 2 months) were representa the variety Moudouma (Table 1). The general decrease in tive of the maximum number usually observed 2 months leaf area was found to depend on the plant itself. Com after the beginning of gradation in Congo (Le Rü et al., pared to Schulthess et al. 's (1991) work on only one variety 1991b). The artificial infestation is done on the second and of cassava (TMS 30572), our study shows that although third leaves from the apex with young neonate larvae. the infestation of P. manihoti is characterized by a de They attach themselves to the plant within the first 48 crease in leaf area, it can be variable depending on the hours, resulting in a homogeneous infestation of plants by variety, and ranges between 7% and 56%. When plant P. manihoti. We then count and readjust their number to growth was analysed in detail-for instance by taking into 100 (by removing or adding mealybugs of the same stage) account the average number of leaves, number and aver during ail experiments, in order to have an identical num age length of internodes-we then found that the develop ber on ail infested plants. Finally, when mealybugs reach ment of plants was affected only to a small extent by the L4 stage with ovisacs, we remove the ovisacs each mealybug infestation (Table 1). When infestation modified week to prevent new larvae from hatching and to keep the plant growth significantiy, it decreased either the lifespan total number constant. of leaves (30 M7) or their rate of development (Talinum) or The following parameters were determined at time To both (3 M8). Our results do not suggest a general pattern and T1 on control and experimental plants for each host for the effect of mealybug infestation on the growth of plant: the average number of leaves, the average number cassava.