Studies on the Physiology of Host-parasite Relationship in Orobanche. I. Respiratory Metabolism of Host and Parasite

The respiratory rate of the roots of mustard (Brassica cam-pestris L.) and tomato (Lycopersicum esculentum Mill.) serving as hosts for the total root parasites Orobanche aegyptiaca Pers. and O.cernua Loefll. was measured using Warburg manometric technique. At the same time determinations were made of the respiration of the apical, basal and root regions of the parasites. The effects of sodium fluoride, malonic acid, sodium azide and DNP (2,4-dinitrophenol) on the rate of respiration of the host roots as well as of the parasites were studied. The Orobanche infection results in a marked increase in the respiratory rate near the host-parasite contact region. The damaging effect of infection seems to be due mainly to a continuous flow of water, minerals and metabolites from host to parasite. The haustorial invasion creates an obstruction in the translocation of metabolites. The respiration rate is lower in Orobanche than in the host, which might be related to its slower growth rate, inefficient oxidative processes and an escaping of certain energy-requiring interconversion processes. Roots of O. aegyptiaca are more well-developed and have higher rate of respiration. They can absorb more water and minerals from the soil. This fact might be connected with the specificity of the two species. NaF and malonic acid inhibit the respiration to a similar extent in healthy and infected roots. This indicates that the pathway of respiration does not change materially after infection. The EMP and Krebs cycle seem to operate at a lower intensity in Orobanche, which is proved by the lower inhibition of the respiration as compared to in the host. Azide causes a stronger reduction of the respiration in infected than in healthy roots. It would imply that the infection stimulates the activity of metal containing oxidases. The weaker inhibition of the respiration in Orobanche tissues indicates a mediation of other enzymes in the oxidation processes than in the host. The respiration is less stimulated by DNP in infected than in healthy roots. Contrary to the general effect of DNP, this substance decreases the O₂ uptake in the parasite tissues. This fact may be explained by the occurrence of exceptionally high amounts of endogenous phenolic compounds and an insufficient production of ATP in the parasite.     

Studies on the Physiology of Host-parasite Relationship in Orobanche. III. Carbohydrate and Nitrogen Metabolism of Host and Parasite

Determination of different carbohydrate and nitrogen fractions was made in tomato (Lycopersicum esculentum Mill.) and mustard (Brassica campestris L.) serving as hosts for Orobanche cernua and O. aegyptiaca respectively. Shoots of Orobanche were also subjected to such analyses. Infection raised the level of total reducing and total sugar in the host with a simultaneous decrease in the level of acid-hydrolyzable and total carbohydrates in the constituent organs of infected hosts. This has been explained to be due to predominance of hydrolytic activity in the infected host. Infection also brought about a depression in the proportion of sucrose to the pool of total sugar in the host. This was also possibly due to predominance of hydrolytic processes and retardation in the synthetic processes. Higher concentration of acid-hydrolyzable and total carbohydrates in Orobanche than in the host indicated a high demand for sugars by the parasite. The insignificant differences between the relative proportions of different nitrogen fractions to the pool of total nitrogen in healthy and infected hosts indicated that nitrogen metabolism was not deranged in any way due to infection. Orobanche always had a lower concentration of total soluble and total nitrogen than the host root.     

Mineral Nitrogen in Plant Physiology and Plant Nutrition

Nitrogen (N) nutrition enhances metabolic processes that influences the physicochemical environment at the soil-root interface, modifies rhizosphere conditions, interferes with the uptake of cations and anions, and enhances or represses the activity of several enzyme systems. Also, it affects growth patterns, protein content, and protein quality of seeds.

Ammonium (NH₄)-N nutrition increases anion uptake, free amino-N/protein ratios, and acidity of root free space; it reduces carbohydrate levels in plant tissues. NO₃-N nutrition results in higher cation uptake, higher carbohydrate content in tissues, and alkalinization of root free space. N-Assimilation interferes with the allocation of dry matter and energy, which causes different growth rates of plant parts.

In this article we review the effects of mineral-N nutrition on uptake of cations and anions, activity of enzymes, growth patterns of roots and shoots, and water use efficiency, protein content, and protein quality of seeds.     

Studies in the Nutrition of Apple Rootstocks: III. Effects of Level of Magnesium Nutrition on Growth, Form and Mineral Composition

One-year rooted shoots of apple M. VII rootstock were grownfor a single season by spraying their roots continuously withnutrient solutions containing <3, 10, 45, and 135 ppm Mg.These treatments respectively produced plants with a severeMg deficiency (Mg₍₀₎), a mild deficiency (Mg₍₁₎), normal foliageand growth (Mg₍₂₎), and a high level of Mg (Mg₍₃₎). The effectof Mg nutrition on the leaves can explain most of the observedeffects on vegetative growth. Depression of growth of Mg₍₁₎)plants was not detected until late in the season although necrosis,typical of Mg deficiency, occurred on the lower leaves by mid-July.This loss of photosynthetic area of leaf was to a large extentcompensated for by increased efficiency of the remaining greenleaves; as the deficiency symptoms progressed, the loss of leaf,and therefore of carbohydrate production, predominated and totalgrowth was affected. The form of the plant was little altered. Symptoms were apparent on leaves of severely deficient Mg₍₀₎plants by late June, and reductions in growth and modificationsin form by August. The poor root systems and thin shoots arelargely attributed to loss of leaf rather than to the low Mgconcentration in these organs. No symptoms of toxicity developed on the Mg₍₃₎ plants althoughby the end of the season the total dry weights were less thanthose of the Mg₍₂₎ controls, and the plants also produced thickershoots. Plant dry weight, and more especially Mg uptake, increasedfrom Mg₍₀₎ to Mg₍₂₎ but above this level the gradient of Mguptake in relation to supply was less steep. The excess Mg takenup at the Mg(j) level led to the relatively lower weights ofthese plants. Much of the excess Mg accumulated in the leaves,especially in the older ones. Mg moved from, or accumulatedin, the lower leaves of the shoot according to the level ofMg supplied to the plants. Thus the shoot apex tended towardsnormality both in the concentration of Mg and in its growth. In general, growth of the plant, and of individual componentparts, governed the uptake of elements other than Mg, althoughmore K, Ca, and Mn and less Fe per unit of plant dry weightaccumulated at the lower levels of Mg than at the higher levels.     

Physiological Studies in Plant Nutrition: XVI. The Mineral Nutrition of Bracken

Part I of this paper describes culture methods for the productionof large numbers of bracken sporelings of uniform size. Observationson the growth of the bracken plant and results of experimentson the mineral nutrition of the prothallial and sporophyte generationsare presented. The presentation of further growth data and afull analysis of the results is deferred to the second partof this paper.     

Physiological Studies in Plant Nutrition: XVI. The Mineral Nutrition of Bracken

1. The nutritional requirements of the bracken sporophyte wereexamined in a factorial combination of 3 potassium levels x2phosphorus levels x3 solution types, in which the cations weremainly Na, Ca, or NH₄.
2. The effects of nitrogen and phosphorusdeficiency and of shadingunder conditions of high and low potassiumsupply were alsoexamined.
3. Leaf area, total dry weight, andnet assimilation rates aremuch depressed by lack of eitherK or P.
4. Water contents of leaves and rhizomes are generallyincreasedby lack of K when Na is present in the culture mediumbut notwhen Ca is in excess.Phosphorus causes diminished succulence.
5. Starch content increases as phosphorus supply is lowered.Withincreasing doses of K, starch content falls in the highcalciumsolution, while increasing in the other two solutiontypes.
6. Reduction of light intensity is shown to have a beneficialeffectunder conditions of K deficiency.
7. Analysis revealscomplex interactions between the various factors,and possibletoxic effect of Na, NH₄, and excess P. An estimateof the relativeimportance of net assimilation rate, leaf numberand area perleaf in determining total plant size has been calculatedforthe different nutrient treatments. Comparisons are madewithprevious results for barley and flax.

     

Studies in the Nutrition of Apple Rootstocks: IV. The Effect on Growth and Mineral Coomposition of Supplying Magnesium through the Leaves

One-year rooted shoots of M.VII apple rootstock were grown fora single season by spraying their roots continuously with nutrientsolutions containing either < 3 ppmMg(Mg₍₀₎) or 45 ppm(Mg₍₀₎)to give, respectively, potentially very deficient or healthyplants. The new shoots of half the plants in each of these treatmentswere dipped periodically in a 2 per cent solution of MgSO₄.₇H₂O plus ‘wetter’. Mg₍₀₎ undipped plants developed severe symptoms of Mg deficiency,growth was poor, and the shoot/root dry-weight increment ratiowas high; none of these characteristics was found in Mg₍₀₎ dippedplants, whose growth was not appreciably less than that of Mg₍₂₎undipped controls. There was little translocation of Mg from leaves to roots: theconcentration of Mg in roots of Mg(0) dipped plants was as lowas that of the undipped. The large accumulation of Mn and, toa less extent, of Fe in Mg₍₀₎ dipped roots was not apparentlydetrimental to growth. Growth of Mg⁽²⁾ dipped plants was similar to that of the undipped. Dipping had little effect on the chemical composition of leaves,except to raise the concentration of Mg.     

Studies in the Nutrition of Apple Rootstocks The Effect on Growth and Mineral Composition of Supplying Iron Through the Leaves

Apple rootstocks were grown with either 0.02 ppm Fe (Fe₀) or5 ppm (Fe₃), to give very chlorotic or dark-green plants. Toinvestigate whether iron can be supplied through leaves insteadof roots the shoots of half the plants in each treatment weredipped periodically in solutions of iron. This prevented chlorosisin Fe₀ plants and increased their growth, which did not, however,equal that of Fe₃ plants supplied with iron through the roots.Growth of Fe₃ plants was reduced by dipping. Iron was not translocated from leaves to roots, although theconcentration in leaves was greatly increased by dipping. Dippingreduced the amount of manganese in Fe₀ roots to one-quarterof that in roots of undipped Fe₀ plants. Effects of treatmentson nitrogen, potassium, calcium, magnesium, and copper levelsare also described.     

The Regulation of the Water Potential Gradient in the Host and Parasite Relationship betweenSorghum bicolorandStriga hermonthica

A glasshouse experiment was carried out to investigate the factorscontrolling the abstraction of xylem fluid from its host bythe parasiteStriga hermonthica(Scrophulariaceae).Strigahad amean daily transpiration rate far exceeding that of its hostsorghum (Sorghum bicolor), with infestation byStrigaalso shownto lower the transpiration rate of the host. Stopping the host'stranspiration was shown to decrease the transpiration rate ofthe parasite. Stopping the parasite's transpiration only gavean initial increase in the host's transpiration rate which wasnot sustained. The parasite had a lower water potential thanits host, values being -0.42 MPa and -0.23 MPa, respectively,and an accompanying higher osmotic pressure of 0.68 MPa against0.51 MPa for sorghum. Modifying the water potential gradientby bagging both partners together showed that the differentialin osmotic pressure and water potential was largely maintainedby the parasite's higher rate of transpiration. A favourablewater potential gradient towards the parasite still existedfollowing the cessation of transpiration, this being generatedby the haustorial resistance to hydraulic conductivity whichwas found to be some 1.5–4.5 times greater than that offeredby the parasite shoot. Both the high rate of transpiration andthe increased resistance across the haustoria would appear tobe necessary means to facilitate the diversion of host resourcesto the parasite.Copyright 1997 Annals of Botany Company Striga hermonthica; sorghum; water relations; haustorium; root parasite     

Studies on Orobanche hederae Physiology: Pigments and CO2 Fixation

In order to investigate some aspects of Orobanche hederae physiology in relation to its parasitism, the pigment composition and the ¹⁴CO₂ incorporation, both in light and in dark, were studied. By means of various chromatographic techniques, it was shown that chlorophyll is probably quite absent and that the carotenoids are very largely distrihuted and consist almost exclusively of xanthophylls. Flavochrome, flavoxanthin, and a pigment resembling neoxanthin were found to be the major components; lutein-5,6-epoxide, taraxanthin, and traces of β-carotene and α-carotene-5,6-epoxide were also present. CO₂ incorporation is completely of heterotrophic type, being in no way stimulated by the light; the major radioactivity was detected in the fractions containing amino acids and organic acids.     

The Effect of Mineral Nutrition on the Distribution of Growth in Lolium perenne

Ramets of Lolium perenne from the same genotypes were grownin nutrient solutions of two concentrations. Additional mineralnutrition increased the rate of production of new tillers, therate of increase in size of tillers and the rate of productionof new roots, but not the rate of growth of individual mainroots. The genotypes differed considerably from each other intheir rates of growth, especially of shoot growth. When thegrowth-rate of the same population of genotypes was alteredby changing the environmental conditions it was found that thelower the mean rate of shoot growth, the greater its coefficientof variation. Variation in the rate of shoot growth, due tomineral nutrition and genetic constitution of the plants growingin a dilute nutrient solution, was accompanied by a similarbut smaller variation in the number of roots.