Anatomical responses of root tips to boron deficiency II. Effect of boron deficiency on the cellular growth and development in root tips

Changes induced in the tissue structure and the cellular patterns of young tomato root tips by the absence of boron in the nutrient solution were investigated.

Boron deficiency caused primarily the inhibition of cell division and cell elongation in root apices, and the cells of boron-deficient root tips were fully vacuolated. The cell wall in the apical region was thickened by boron deficiency and the intercellular spaces insufficiently developed.

Boron deficiency also caused the radial enlargement of cortical cells, especially of endodermis, but this enlargement was not accompanied by an increase in water imbibition. In the advanced stage of boron deficiency, the disintegration of tissue structure had occurred.

Primordia of lateral roots arose closely in root apices. Maturation of the vascular system, especially of the primary xylem, was exasperated abnormally, and frequently there occurred a differentiation of cambial layers close to the apical initials.

Anatomical effects of boron deficiency appeared particularly in the root apex and not clearly in the region of successive maturation. The results are discussed with regards to the role of boron in cellular growth at apical growing points.

The results are discussed with regards to the role of boron in cellular growth at apical growing points.     

缺硼豌豆植株侧芽生长机理的研究

用营养液培养的方法对供硼与不供硼（缺硼）条件下豌豆植株的生长及侧芽发育状况进行了研究。结果表明,缺硼明显抑制豌豆植株生长,根系受抑先于地上部。缺硼１０天后可见侧芽长出,顶端优势不明显。用间接酶联免疫法（ＥＬＩＳＡ）测定了内源生长素（ＩＡＡ）的细胞分裂素（Ｚ／ＺＲ）水平的动态变化,发现缺硼植株顶芽中ＩＡＡ水平高于对照,顶芽的可扩散ＩＡＡ水平低于对照,节中ＩＡＡ水平降低;缺硼初期（３～６天）,根系和节中Ｚ／ＺＲ水平均高于正常供硼植株。认为缺硼豌豆植株节中的低ＩＡＡ水平和高Ｚ／ＺＲ水平是诱导侧芽发育的一个重要因素     

Symptome von Bormangel und Borüberschuß bei Hibiscus esculentus

Boron deficiency and Boron toxicity in Hibiscus esculentus The development of symptoms in Okra under conditions of B-deficiency and B-toxicity was observed. It was compared if the symptoms correspond with the B-contents and the growth of the plants. Symptoms of Boron deficiency in Okra are the same as in the other annual dicotyledons. The first visible microscopical symptom is a disturbance in the cambium region. Characteristic is an enlargement of the cambium. New divided cells do not differentiate. Visual symptoms are breaking of stems and petioles, thickening of these organs, cracking, dying of the tips and falling of buds. With a periodic supply of Boron correlations between Boron content and deficiency symptoms were not found. In the B-toxicity range small yellow chlorosis developed on the margins of the older leaves. These symptoms are not to be used for diagnosis. Boron contents higher than 72 μg B/g DM gave a better measure for B-toxicity. Growth and development of symptoms were in agreement with the high Boron contents.     

Makro-und Mikrosymptome des Bormangels bei Sonnenblumen,Chinakohl und Mais

Macro- und microsymptoms of boron deficiency of sunflowers, Chinese cabbage and com Boron deficiency leads to the same tissue aberrations as well in mono-as in dicotyledons; disturbances in the differentiation of vascular bundle-and mesophyll cells. Furthermore dicotyledons show an increased activity in cell division in the fascicular and interfascicular cambiums even before the apical meristem of the main shoot dies. Mainly phloem parenchyma cells are produced. In severe long persistent deficiency the result of this is an extention of the phloem, expecially in the lower internodes. Xylem cells (particularly vessels and tracheids) which are produced by the cambium in a lower quantity remain thin walled and more or less parenchymatic.     

γ射线辐照对大蒜和洋葱生长点细胞及超微结构的影响

电镜观察表明,γ射线2～30 krad辐照大蒜和洋葱,能造成生长点细胞及超微结构的不同程度损伤。如细胞外形畸变、细胞壁加厚、质壁分离、原生质破坏呈电子透明,核破碎,染色质变粗变短,微结构变形,甚至膜破裂等。其损伤程度随辐照剂量的增加而趋严重。不同微结构的抗辐照能力不同,其中液泡的辐射敏感性最强,核仁和线粒体最抗辐照。辐照后的细胞和超微结构一般在萌动期才表现出明显的形态损伤、细胞分裂受阻,因此抑制了大蒜和洋葱的发芽。     

The physiological role of boron in plants

Research on the role of boron in plant metabolism is reviewed and evaluated. Absence of the element is shown to affect nucleic acid and carbohydrate metabolism, and the relationship between boron, auxin and phenolic compounds is discussed. It is concluded that the presence of boron is essential to maintain the structural integrity of plant membranes, and that many symptoms of boron deficiency in plants are secondary effects caused by changes in membrane permeability.     

Der Einfluß von Dicyandiamid und N-Serve in Verbindung mit Ammoniumsulfat als N-Dünger auf die Nitrat- und Oxalsäuregehalte von Spinat (Spinacia oleracea)

Occurence of boron deficiency in Citrus aurantium L. (Bitter orange) at the Caspian Sea (Iran) In a Bitter orange plantation near the Caspian sea over a period of two years different amounts of boron were applied either as borax to the soil or as Solubor to the leaves. Without boron supply the fruits were misshapen, had a rough and thick peel and often brown spots in the pulp. Supply of 5 – 20 g B per tree prevented the occurence of these symptoms and increased both the fruit size and the proportion of the pulp. Without boron supply on fresh weight basis the proportion of pulp/peel was 1 : 1, 6 – 2, 0 as compared to 1 : 0, 6 – 0, 7 with boron supply. The effect of the different boron supply was well reflected in the boron content of the leaves and especially of the fruits. Borax increased the boron content of the plants more than Solubor. Conzidering both the low content of available boron in most of the soils of the Citrus area near the Caspian sea and the special ecological conditions (high pH values and low water content of the soils during fruit development) boron deficiency can be expected. The results of the experiment with Bitter oranges confirm that boron fertilizing is an important factor for plant production in this area.     

Gewebekulturen unter Bormangel

Tissue culture under conditions of boron deficiency Cambium tissue from sunflowers and carrots were cultured in nutrient solution according to Murashige and Skoog with graduated supplies of boron. A light-coloured and well growing callus was formed only in the presence of boron. Explants on a boron-deficient medium agglutinated at the edges, had only small areas of cell division and revealed appositions on the walls of some cells. Deficient tissues also became noticeably darker. Callus grown on normal medium and then transferred to boron-deficient medium slowed down in growth rate – at the latest after 4 weeks – and also turned dark. In contrast to normal calli, the deficient cultures could not be easily separated in water into individual cells or small cell groups. Moreover, the cells were smaller on the average and often revealed grainy contents and (in the case of carrots) accumulation of plasma at the ends of the cells. A large number of deficient cells were plasmolyzed. An accumulation of undifferentiated cells in the cambium region is particularly striking in dicotyledons under conditions of boron deficiency. If this symptom should be the result of cell division activity, then isolated cambium should react to boron deficiency with intensive cell division. Contrary to expectation, isolated cambium of sunflowers and carrots in tissue culture did not reveal a promotion of cell division under conditions of boron deficiency. These results indicate that the symptom of “increased cambium growth” in intact dicotyledons is due to a secondary effect of boron deficiency.     

缺硼对脐橙幼苗硼分配及叶片细胞壁组分硼含量的影响

【目的】 硼在维持细胞壁正常结构方面具有重要的作用,前期结果证实缺硼严重的脐橙叶片细胞壁结构改变程度也更大,但这种变化与细胞壁组分中硼的含量变化是否有关尚不清楚。本研究通过分析缺硼对脐橙幼苗各部分硼分配及叶片细胞壁组分硼含量的影响,明确缺硼症状表现及细胞壁结构变化程度与细胞壁各组分中硼含量变化之间的关系。 【方法】 以纽荷尔脐橙幼苗为试材,利用营养液培养方法进行缺硼处理,测定根、砧木茎、接穗茎、上部叶、下部叶、叶片细胞壁以及细胞壁各组分硼含量的变化情况。 【结果】 缺硼处理9周后上部叶出现叶片卷曲及叶片失绿等症状,而下部叶没有出现任何可见的症状。缺硼处理的脐橙幼苗各部位硼含量和硼吸收量均显著降低,缺硼降低了硼向地上部的相对分配比例且上部叶受到的影响程度更大。在硼正常供应条件下,上部叶和下部叶游离态硼、原生质体硼和细胞壁硼的含量和相对分配比例没有显著差异,说明硼在不同类型脐橙叶片细胞各组分中的分配是相对稳定的。缺硼后水溶性硼 (包括游离态硼和原生质体硼) 在脐橙上部叶和下部叶中都降到极低的水平,尤其是原生质体硼百分含量下部叶甚至是低于上部叶的。缺硼后细胞壁硼占总硼的比例则由22%左右增加到80%以上。与叶片中硼含量的变化趋势一致,缺硼以后虽然上部叶和下部叶细胞壁硼含量都显著降低,但上部叶降低的程度远大于下部叶。进一步分析细胞壁组分硼含量变化,发现缺硼显著降低了上部叶细胞壁中离子结合态果胶硼含量而对下部叶的无明显影响,其他组分硼含量的变化趋势下部叶和上部叶一致。 【结论】 原生质体硼含量的高低并不是决定缺硼症状的主要因素,离子结合态果胶与硼的结合能力对缺硼条件下细胞壁的结构及缺硼症状表现起着至关重要的作用。      

缺硼槟榔幼苗的生理反应和根系发育特征

  【目的】  探究缺硼对槟榔幼苗生理特征和根系形态的影响,以期为槟榔缺硼（B）诊断提供理论依据。  【方法】  以‘热研1号’槟榔幼苗为材料进行了砂培试验。设置营养液中不加B (B0,0 μmol/L) 和添加常规硼 (B50,50 μmol/L) 两个浓度处理。生长3个月后,测定了槟榔幼苗生物量、株高、硼含量、叶片糖类物质以及抗氧化酶活性、丙二醛含量、光合速率,观察了不同处理下根尖及根尖细胞的形态。  【结果】  与B50处理相比,B0处理显著降低了槟榔幼苗株高、地下部鲜重和干重、总鲜重、地上部和地下部硼含量;B0处理槟榔叶片蔗糖和淀粉含量下降,可溶性总糖含量无显著变化,叶片光合速率降低,且MDA含量、POD活性显著升高。在B0条件下,槟榔根系变短,根尖明显膨大,细胞壁明显增厚,且内壁上积累了大量的颗粒物,根系活力显著降低。  【结论】  缺硼导致槟榔幼苗根尖解剖结构受到破坏,养分吸收运转能力降低,叶片抗氧化系统受到损伤,光合能力下降,最终抑制植株生长。     

Differential Changes in Cell-Wall Content and Boron and Calcium Concentration in Newhall Navel Orange Grafted on Two Rootstocks Differing in Boron-Deficiency Responses

The study aimed to determine if the variability of the effect of boron (B) deficiency on Newhall navel orange grafted on trifoliate orange (deficient-B-sensitive) and on citrange (deficient-B-tolerant) can be explained on the basis of changes in cell-wall content and in cell-wall B and calcium (Ca) concentrations. The plants were cultured in the nutrient solution (with or without B) for 65 days. Boron deficiency increased the cell-wall content in old leaves (leaves from last season) of trifoliate orange but had no impact on citrange. Boron deficiency did not reduce B concentration in cell walls of old leaves of citrange-grafted plants but increased their Ca concentration. For trifoliate-orange-grafted plants, however, B deficiency decreased the B concentration in cell walls of old leaves and did not increase their Ca concentration. The changes of B and Ca concentrations between with and without B supply in pectin were in good agreement with the changes of those in cell walls. The relatively greater ability of citrange-grafted plants to maintain B and Ca in the cell wall may contribute to their tolerance to low external B.     

Wirkung hoher Phosphatdüngung auf die Wachstumsrate,den Zinkgehalt und das P/Zn-Verhältnis in Weinreben (Vitis vinifera)

Effect of high phosphate fertilization on growth rate, zinc content and P/Zn ratio in grapevines (Vitis vinifera) In a 2 years' pot experiment with a soil low in available Zn, high doses of phosphate fertilizer induced Zn deficiency in grapevines (var. ?Riesling”? on 5 C root stock). This negative effect of phosphate fertilization could almost be prevented by simultaneous application of Zn to the soil. The negative effect of high phosphate fertilization on the Zn status of the soil was well reflected in both the content of available Zn as well as in the diffusion rate of Zn (⁶⁵Zn) in the soil. In young leaves of grapevines, Zn deficiency can be expected at Zn contents lower than 20 μg/g DM and P/Zn ratios larger than 170. These values, however, cannot be generalized as external factors modify them considerably. This latter holds true for the P/Zn ratio in particular. The most obvious Zn deficiency symptom was restricted elongation of the shoot due to shorter internodes. Typical for severe Zn deficiency were smaller leaves with intervenal chlorosis or even necrosis and the disappearance of the normally deeply lobed shape of the leaf blade. As a result of the growth inhibition of the shoot apex, growth of axillary buds had been enhanced and this led to a bushy shape of the Zn-deficient plants.     

Makrosymptome des Bor-Mangels an Mentha piperita L.

Macrosymptoms of boron deficiency in Mentha piperita L. The influence of boron deficiency on growth and development of Mentha piperita L. – var. Mitcham – was studied under controlled experimental conditions in solution culture. Acute boron deficiency depressed growth of stems and roots. Furthermore the following symptoms – well known as boron deficiency symptoms – occured in peppermint: restricted growth of internodes, leaf distortion, simplification of leaf form, dropping of parts of organs, bursting of cortex tissue, dis – colouration, chlorosis and necrosis. The typical symptom in dicots – death of growing point – could not be observed in peppermint. The visible symptoms of boron deficiency in peppermint are typical for that micronutrient. They can be helpful in diagnosing and curing that deficiency.     

Symptome von Mangan-Überschuß bei Bohnen (Phaseolus vulgaris)

Symptoms of manganese toxicity in beans (Phaseolus vulgaris L) In water culture experiments with bush bean plants (Phaseolus vulgaris L. var. ?Red Kidney”?) the macro- and micro-symptoms of manganese toxicity had been studied. Typical symptoms of Mn toxicity are small distinct blackish-brown spots, first at the older leaves and at more severe toxicity also at the petals and stems. Using the autoradiographic technique on ⁵⁴Mn-distribution and comparing this with corresponding light microscopic studies it could be demonstrated that the blackish-brown spots in the leaves resemble Mn-precipitations. These precipitations are Mn-compounds of higher oxidated form and are primarily localized in the walls of cells in vicinity of the vessels. The extend to which these precipitations occur is not only a function of the Mn-content of the leaves but also of their age. At comparable Mn-content the extend of the precipitation increases with the leaf age. As secondary symptom of Mn-toxicity under certain conditions (presence of silicon e.g.) induced Fe deficiency (chlorosis of the youngest leaves) and Ca deficiency (?crinkle leaf”?) may occur. It is assumed that the process of Mn precipitation within the leaves is enzymatically regulated. The severe reduction in the growth of leaf blades at Mn toxicity is presumably related to an interference with the growth hormon balance.     

PHYSIOLOGICAL AND BIOCHEMICAL CHANGES IN CATHARANTHUS ROSEUS L. IN RESPONSE TO BORON NUTRITION

Catharanthus roseus L., a medicinally important plant was grown till maturity at varying levels of boron (0.033, 0.066, 0.33 and 3.3 mg B L^?1) supply. Optimum yield was observed in plants receiving 0.33 mg B L^?1. Plants receiving deficient boron showed growth reduction and visual symptoms such as chlorosis and cupping of young emerging leaves and apical tip necrosis. The number and size of the flowers, pods and seeds formed and the pollen viability of the boron deficient plants was markedly reduced. The threshold values for deficiency and toxicity were 57 and 79 μg B g^?1 dry weight for vegetative growth and 60.4 and 68.9 μg B g^?1 dry weight for reproductive growth, respectively, and is reported for the first time in periwinkle. An increase was observed in reducing and non-reducing sugars and in activity of acid phosphatase and ribonuclease in boron stressed plants.     

Factors affecting shoot and root apical meristem tissue culture of Thai supersweet corn (Zea mays var. rugosa)

During last recent years, in vitro propagation technic is widely used to produce plants with desirable traits. This experiment was conducted to produce an ideal protocol for in vitro propagation of Thai supersweet corn by using shoot apical meristem (SAM) and root apical meristem (RAM) as explants. Four-day-old germinating seedlings were used as the experimental materials on culture media supplemented with a range of auxin, kinetin, and carbohydrates. The primary establishment for SAM showed the highest percentage of survival (80%) while RAM showed the highest survival (67%) and in Murashige and Skoog (MS) media supplemented. Upon acclimatization, regenerated plantlets from shoot showed the highest survival rate (12%) with the production of 21 plantlets; however, the survival rate of plantlets from root was only 20% with the production of 9 plantlets. The efficient and economic protocol that is produced in this study can be applied as an alternative to conventional propagation method for the large-scale production of Thai supersweet corn throughout the year.     

Depressed growth of non‐chlorotic vine grown in calcareous soil is an iron deficiency symptom prior to leaf chlorosis

The development of iron deficiency symptoms (growth depression and yellowing of the youngest leaves) and the distribution of iron between roots and leaves were investigated in different vine cultivars (Silvaner, Riparia 1G and SO4) grown in calcareous soils. As a control treatment all cultivars were also grown in an acidic soil. Only the cultivars Silvaner and Riparia 1G showed yellowing of the youngest leaves under calcareous soil conditions at the end of the cultivation period. All cultivars including SO4 showed severe shoot growth depression, by 50 % and higher, before yellowing started or without leaf yellowing in the cultivar SO4. Depression of shoot growth occurred independently from that of root growth. In a further treatment the effect of Fe‐EDDHA spraying onto the shoot growth of the cultivar Silvaner after cultivation in calcareous soil was investigated. Prior to Fe application plants were non‐chlorotic, but showed pronounced shoot growth depression. Spraying led to a significant increase in shoot length, though leaf growth was not increased. Accordingly, depression of shoot growth of non‐chlorotic plants under calcareous soil conditions and with ample supply of nutrients and water has been evidenced to be at least partly an iron deficiency symptom. We suggest that plant growth only partially recovered because of dramatic apoplastic leaf Fe inactivation and/ or a high apoplastic pH which may directly impair growth. Since growth was impaired before the youngest leaves showed chlorosis we assume that meristematic growth is more sensitively affected by Fe deficiency than is chlorophyll synthesis and chloroplast development. In spite of high Fe concentrations in roots and leaves of the vines grown in calcareous soils plants suffered from Fe deficiency. The finding of high Fe concentrations also in young, but growth retarded green leaves is a further indication that iron deficiency chlorosis in calcareous soils is caused by primary leaf Fe inactivation. However, in future, only a rigorous study of the dynamic changes of iron and chlorophyll concentration, leaf growth and apoplastic pH at the cellular level during leaf development and yellowing will provide causal insights between leaf iron inactivation, growth depression, and leaf chlorosis.<?show $6#>     

Response of Hibiscus Rosa‐Sinensis L. to varying levels of potassium fertilization: Growth,gas exchange and mineral element concentration

Little is known about the effect of varying levels of potassium (K) on the mineral element concentration, growth, and gas exchange, characteristics of woody ornamental plants. The commercially important woody ornamental species Hibiscus rosa‐sinensis L. cv. Leprechaun was evaluated for K response in a series of three experiments with full strength Hoagland's nutrient solution, which supplied 0 to 10 mM K. Plants grown with 4 mM K in nutrient solution (2.4% leaf tissue K) had the greatest shoot growth and root extension. Gas exchange rates (net photosynthesis, transpiration, and stomatal conductance) were also highest at 4 mM K compared to the control (0 mM K /0.6% leaf tissue K), 0.2, 2.0 and 10 mM K treatments. The application of 4 mM K increased net photosynthesis and tranpiration by 2.1 fold and stomatal conductance by 4.5 fold over 0 mM K controls. Increasing K in nutrient solution correlated positively with tissue K, manganese (Mn), and zinc (Zn), but negatively with nitrogen (N), phosphorus (P), calcium (Ca), and magnesium (Mg). There was a stronger sink for K in yonger leaves (the first to fourth fully expanded leaf from the shoot apex) which had higher K concentration than older leaves (the eighth to twelfth fully expanded leaf from the shoot apex). However, with increasing K in nutrient solution, K concentration in leaf tissue increased regardless of leaf age, and the difference between the younger and older leaf was constant. Daily application of 10 mM K resulted in 6.9% leaf tissue K and caused a decrease in plant total dry matter, net photosynthesis, compared to 4 mM K treated plants. However, these parameters remained higher in 10 mM K plants, which retained high ornamental quality than in 0 mM controls. Plants fertilized with 10 mM K, had the highest leaf tissue K and Zn, but lowest P, Ca, Mg, iron (Fe), copper (Cu) and boron (B). Nevertheless, the 10 mM K treated plants exhibited no morphological differences or deficiency symptoms; rather those plants had similar vegetative vigor and flower bud formation rate as those at 4 mM K.     

Iron deficiency and zinc toxicity in soybean grown in nutrient solution with different levels of sulfur

A typical symptom of iron (Fe) deficiency in plants is yellowing or chlorosis of leaves. Heavy metal toxicity, including that of zinc (Zn), is often also expressed by chlorosis and may be called Fe chlorosis. Iron deficiency and Zn toxicity were evaluated in soybean (Glycine max [L.] Merr.) at two levels each of Zn (0.8 and 40 μM), Fe (0 and 20 μM), and sulfur (S) (0.02 and 20 mM). Reduction in dry matter yield and leaf chlorosis were observed in plants grown under the high level of Zn (toxic level), as well as in the absence of Fe. Zinc toxicity, lack of Fe, and the combination of these conditions reduced dry matter yield to the same extent when compared to the yield of the control plants. The symptoms of Zn toxicity were chlorosis in the trifoliate leaves and a lack of change in the orientation of unifoliate leaves when exposed to light. The main symptoms of Fe deficiency were chlorosis in the whole shoot and brown spots and flaccid areas in the leaves. The latter symptom did not appear in plants grown with Fe but under Zn toxicity. It seems that Fe deficiency is a major factor impairing the growth of plants exposed to high levels of Zn. Under Zn toxicity, Fe and Zn translocation from roots to shoots increased as the S supply to the plants was increased.     

The influence of Al on the metabolism of spruce needles

Aluminium toxicity as a consequence of increasing soil acidity is discussed as a primary factor in forest decline in middle Europe. Ecologically relevant concentrations of Al disturbed root systems of hydroponically grown spruce (Picea abies Karst.) seedlings. The needles, with a much lower Al content in comparison to the roots, were also affected by Al at these external concentrations. In this investigation needle chlorosis as the visible symptom of Al toxicity, was accompanied by decreasing pigment concentrations and photosynthetic rates as well as increasing starch levels and transpiration rates. The magnesium concentrations of roots and needles were lowered by treatment with Al. By feeding the needles directly with Mg, it was possible to differentiate between direct and indirect effects of Al on needle metabolism. Magnesium deficiency was found to drive all the experimental variables except transpiration.