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- 5. 1 THE IMMUNE SYSTEM, FOURTH EDITION CHAPTER 7: THE DEVELOPMENT OF T LYMPHOCYTES © 2015 GARLAND SCIENCE 7–1 In which of the following ways does the developmental pathway of α:β T cells differ from that of B cells? (Select all that apply.) a. Their antigen receptors are derived from gene rearrangement processes. b. When the first chain of the antigen receptor is produced it combines with a surrogate chain. c. Cells bearing self-reactive antigen receptors undergo apoptosis. d. MHC molecules are required to facilitate progression through the developmental pathway. e. T cells do not rearrange their antigen-receptor genes in the bone marrow. 7–2 Which of the following cell-surface glycoproteins is characteristic of stem cells, but stops being expressed when a cell has committed to the T-cell developmental pathway? a. CD2 b. CD3 c. CD25 d. CD34 e. MHC class II. 7–3 Which of the following processes is not dependent on an interaction involving MHC class I or class II molecules? (Select all that apply.) a. positive selection of α:β T cells b. intracellular signaling by pre-T-cell receptors c. negative selection of αβ T cells d. peripheral activation of mature naive T cells e. positive selection of γ:δ T cells. 7–4 If a double-negative thymocyte has just completed a productive β-chain gene rearrangement, which of the following describes the immediate next step in the development of this thymocyte? a. A pre-T-cell receptor is assembled as a superdimer. b. Rearrangement of γ- and δ-chain genes commences. c. Expression levels of RAG-1 and RAG-2 are elevated. d. The linked δ-chain genes are eliminated. e. This cell will inevitably differentiate into a committed γ:δ T cell. 7–5 All of the following cell-surface glycoproteins are expressed by double-negative thymocytes undergoing maturation in the thymus except _____. (Select all that apply.) a. CD2 b. CD5
- 6. 2 c. CD127 (IL-7 receptor) d. CD34 e. CD1A f. CD4. 7–6 _____ is a T-cell-specific adhesion molecule expressed before the expression of a functional T-cell receptor while the thymocytes are still in their double-negative stage of development. a. CD4 b. CD8 c. CD25 d. CD2 e. CD3. 7–7 Which of the following is mismatched: a. double-negative CD3– thymocytes: cortico-medullary junction b. double-negative CD3– thymocytes: subcapsular zone c. double-positive CD3+ thymocytes: cortico-medullary junction d. cortical epithelial cells: subcapsular regions e. dendritic cells: cortico-medullary junction. 7–8 After interaction with thymic stromal cells, _____, a glycoprotein not expressed by the uncommitted progenitor cell is activated in developing thymocytes. (Select all that apply.) a. CD2 b. CD34 c. CD5 d. CD127 (IL-7 receptor) e. CD44. 7–9 Which of the following statements about Notch 1 is correct? (Select all that apply.) a. Notch 1 is expressed on thymic epithelial cells. b. In the absence of Notch 1 expression, T cells can complete their differentiation. c. Notch 1 is to T-cell development as Pax-5 is to B-cell development. d. Notch 1 contains two distinct domains, one of which is proteolytically cleaved and becomes a transcription factor in the nucleus. e. The extracellular domain of Notch 1 must interact with a ligand on thymic epithelium to initiate cleavage and separation of the Notch 1 extracellular and intracellular domains. 7–10 Which of the following is the first stage of T-cell receptor gene rearrangement in α:β T cells? a. Vα→Dα b. Dα →Jα c. Vβ→ Dβ d. Dβ→Jβ e. Vα→Jα.
- 7. 3 7–11 Which of the following is the first T-cell receptor complex containing the β chain to reach the cell surface during the development of T lymphocytes? a. γ:β:CD3 b. β:CD3 c. α:β:CD3 d. β:CD44 e. pTα:β:CD3. 7–12 The T-cell receptor β-chain locus can undergo successive gene rearrangements to rescue unproductive V(D)J rearrangements. A. What aspects of gene segment rearrangement at the TCRβ locus make this possible? B. Can the immunoglobulin heavy-chain locus, which is also composed of V, D, and J segments, undergo successive rearrangements? If not, give the reasons for the difference. 7–13 Indicate which of the following statements is true (T) or false (F). a. __ Immature T cells failing to successfully recombine a β-chain locus die by apoptosis. b. __ Apoptotic T cells are ingested by medullary epithelial cells. c. __ Allelic exclusion of the T-cell receptor α and β chains is effective; therefore, all T cells express only one T-cell receptor on the cell surface. d. __ T-cell receptor rearrangements have many features in common with immunoglobulin rearrangement, including the use of the RAG-1 and RAG-2 genes. e. __ The expression of the pre-T-cell receptor is required in order to halt β-, γ-, and δ-chain rearrangements. 7–14 Genetic deficiencies in all of the following would impair the development of a fully functional T-cell repertoire except a. RAG-1 or RAG-2 b. Notch1 c. Pax-5 d. IL-7 receptor (CD127) e. TAP-1 or TAP-2. 7–15 A. What is Notch1? B. Which cells express the ligand of Notch1? C. How does the interaction between Notch1 and its ligand mediate T-cell development? 7–16 There are many parallels between the development of B cells and T cells. Identify the incorrectly matched counterpart in B cells (left) versus T cells (right). a. VpreBλ5: pTα b. Igα/Igβ:CD3 c. Pax-5: FoxP3 d. multiple κ and λ light-chain gene rearrangements: multiple α-chain gene rearrangements. 7–17 _______ of thymocytes is necessary to produce a T-cell repertoire capable of interacting with self-MHC molecules.
- 8. 4 a. positive selection b. negative selection c. apoptosis d. receptor editing e. isotype switching. 7–18 Which of the following statements are true of a T cell that expresses two α chains (and thus two different T-cell receptors) as a result of ineffective allelic exclusion of the α chain during rearrangement? (Select all that apply.) a. Engaging either of the T-cell receptors on MHC molecules of the thymic epithelium will result in positive selection. b. One of the T-cell receptors will be functional while the other will most probably be non- functional. c. If either T-cell receptor binds strongly to self-peptides presented by self-MHC molecules, the thymocyte will be negatively selected. d. One of the T-cell receptors may be autoreactive but escape negative selection because its peptide antigen is present in tissues other than the thymus. e. Subsequent gene rearrangements may give rise to a γ:δ T-cell receptor. 7–19 Once a thymocyte has productively rearranged a β-chain gene, which of these event(s) can occur subsequently? (Select all that apply.) a. β binds to pTα and is expressed on the cell surface with the CD3 complex and ζ chain. b. Rearrangement of β-, γ-, and δ-chain genes ceases as a result of the suppression of expression of RAG-1 and RAG-2. c. The pre-T cell proliferates and produces a clone of cells all expressing an identical β chain. d. Expression of CD34 and CD2 gives rise to double-positive thymocytes. e. α-, γ-, and δ-chain loci rearrange simultaneously. 7–20 Which of the following statements regarding positive selection is correct? a. All subsets of developing T cells undergo positive selection before export to the peripheral circulation. b. T-cell receptor editing is linked to the process of positive selection. c. Positive selection results in the production of T cells bearing T-cell receptors that have the capacity to interact with all allotypes of MHC class I and class II molecules, and not just those of the individual. d. Positive selection ensures that autoreactive T cells are rendered non-responsive. e. If there is a genetic defect in AIRE, then T-cell development is arrested as positive selection commences. 7–21 Thymocytes that are not positively selected a. undergo genetic reprogramming and differentiate into a different cell type b. are exported to the periphery, where they are phagocytosed by macrophages c. make up about 98% of developing thymocytes and die by apoptosis in the thymic cortex d. are eliminated because of their reactivity with self antigens e. try out different β chains to acquire reactivity with self-MHC molecules.
- 9. 5 7–22 If the process of positive selection did not occur, then a. a condition resembling immune dysregulation, polyendocrinopathy, enteropathy, X- linked syndrome (IPEX) would develop b. a condition resembling autoimmune polyendocrinopathy–candidiasis–ectodermal dystrophy (APECED) would develop c. naive T cells would be unable to undergo differentiation in secondary lymphoid tissues d. malignant transformation would be more likely because of the accumulation of multiple mutations e. only a very small percentage of circulating T lymphocytes would be able to become activated. 7–23 Immediately after positive selection a. the thymocyte reaches maturity and is exported to the periphery b. RAG proteins are degraded and are no longer synthesized c. receptor editing commences to eliminate reactivity against self antigens d. the developing thymocyte acquires a double-negative phenotype e. expression of pTα is repressed. 7–24 Allelic exclusion occurs for all of the following except a. T-cell receptor α genes b. T-cell receptor β genes c. B-cell receptor heavy-chain genes d. B-cell receptor κ-chain genes e. B-cell receptor λ-chain genes. 7–25 A. Explain two ways in which the expression and processing of self antigens in thymic epithelium differs from the expression and processing of self antigens outside the thymus. B. In what way is the thymic situation advantageous for the purposes of negative selection? 7–26 Autoimmune polyendocrinopathy–candidiasis–ectodermal dystrophy (APECED) is caused by a defect in a. cathepsin L b. a transcription factor that regulates tissue-specific gene expression in the thymus c. the production of regulatory CD4 T cells d. FoxP3 e. T-cell receptor gene rearrangement. 7–27 Identify which of the following describes how antigen processing and presentation of self antigens by thymic epithelial cells differs from that of antigen-presenting cells in peripheral tissues. (Select all that apply.) a. Thymic epithelium expresses MHC class I molecules but not MHC class II molecules. b. Thymic epithelium uses cathepsin L for proteolytic degradation of self proteins. c. Thymic epithelium expresses MHC class II molecules but not MHC class I molecules.
- 10. 6 d. Thymic epithelium uses the transcription factor AIRE to activate thymic expression of tissue-specific genes. e. Thymic epithelium expresses transcription repressor protein FoxP3. 7–28 Match the immunodeficiency in Column A with its corresponding cause or consequence in Column B. Column A Column B ___a. IL-7 receptor deficiency 1. absence of functional AIRE ___b. DiGeorge syndrome 2. absence of functional MHC class I or MHC class II molecules ___c. IPEX 3. absence of T cells because of signaling defects by thymic stromal ells ___d. Bare lymphocyte syndrome 4. absence of functional FoxP3 ___e. APECED 5. absence of T cells due to absence of thymus 7–29 All of the following types of protein are processed and presented by macrophages in the thymus except _____ proteins. a. tissue-specific b. soluble proteins from extracellular fluids c. ubiquitous proteins d. proteins made by macrophages e. proteins derived from other cells that macrophages phagocytose. 7–30 Healthy individuals have approximately ____ of CD4 T cells compared with CD8 T cells. a. one quarter the number b. half the number c. equal numbers d. twice the number e. four times the number 7–31 The surrogate light chain operating during pre-B-cell development is made up of VpreB:λ. Its expression with μ on the pre-B-cell surface is an important checkpoint in B-cell maturation. Name the T-cell analog of VpreB:λ5 and discuss how it is functionally similar. 7–32 Double-negative thymocytes initiate rearrangement at the _____ locus (loci) before all other T-cell receptor genes. a. γ and δ b. β c. α and β d. α, γ, and δ e. β, γ, and δ. 7–33 The function of negative selection of thymocytes in the thymus is to eliminate
- 11. 7 a. single-positive thymocytes b. double-positive thymocytes c. alloreactive thymocytes d. autoreactive thymocytes e. apoptotic thymocytes. 7–34 In T cells, allelic exclusion of the α-chain locus is relatively ineffective, resulting in the production of some T cells with two T-cell receptors of differing antigen specificity on their cell surface. A. Will both these receptors have to pass positive selection for the cell to survive? Explain your answer. B. Will both receptors have to pass negative selection for the cell to survive? Explain your answer. C. Is there a potential problem in having T cells with dual specificity surviving these selection processes and being exported to the periphery? 7–35 Mature B cells undergo somatic hypermutation after activation, which, after affinity maturation, results in the production of antibody with a higher affinity for antigen than in the primary antibody response. Suggest some reasons why T cells have not evolved the same capacity. 7–36 MHC class II deficiency is inherited as an autosomal recessive trait and involves a defect in the coordination of transcription factors involved in regulating the expression of all MHC class II genes (HLA-DP, HLA-DQ, and HLA-DR). A. What is the effect of MHC class II deficiency? B. Explain why hypogammaglobulinemia is associated with this deficiency. 7–37 As we age, our thymus shrinks, or atrophies, by a process called involution, yet T-cell immunity is still functional in old age. A. Explain how T-cell numbers in the periphery remain constant in the absence of continual replenishment from the thymus. B. How does this differ from the maintenance of the B-cell repertoire? 7–38 A. What is the role of regulatory CD4 T cells (Treg)? B. How can Treg be distinguished from other non-regulatory CD4 T cells? 7–39 Which of the following statements is correct? a. In adults the mature T-cell repertoire is self-renewing and long-lived and does not require a thymus for the provision of new T cells. b. T cells and B cells are both short-lived cells and require continual replenishment from primary lymphoid organs. c. The human thymus is not fully functional until age 30, at which time it begins to shrink and atrophy. d. In DiGeorge syndrome the bone marrow takes over the function of the thymus and produces mature peripheral T cells.
- 12. 8 e. None of the above statements is correct. 7–40 Individuals with a defective autoimmune regulator gene (AIRE) exhibit a. DiGeorge syndrome b. autoimmune polyendocrinopathy–candidiasis–ectodermal dystrophy (APECED) c. severe combined immunodeficiency (SCID) d. MHC class I deficiency e. MHC class II deficiency. 7–41 Giulia McGettigan was born full term with a malformed jaw, cleft palate, a ventricular septal defect, and hypocalcemia. Within 48 hours of birth she developed muscle tetany, convulsions, tachypnea, and a systolic murmur. A chest X-ray showed an enlarged heart and the absence of a thymic shadow. Blood tests showed severely depleted levels of CD4 and CD8 T cells; B-cell numbers were low but within normal range. Parathyroid hormone was undetectable. Fluorescence in situ hybridization of the buccal mucosa revealed a small deletion in the long arm of chromosome 22. Giulia failed to thrive and battled chronic diarrhea and opportunistic infections, including oral candidiasis and Pneumocystis jirovecii, the latter infection causing her death. Giulia most probably had which of the following immunodeficiency diseases? a. AIDS b. DiGeorge syndrome c. bare lymphocyte syndrome d. chronic granulomatous disease e. hyper IgM syndrome. 7–42 The human thymus begins to degenerate as early as one year after birth. This process is called ______ and is marked by the accumulation of ___ once occupied by thymocytes. a. thymectomy; dendritic cells b. involution; fat c. differentiation; γ:δ T cells d. negative selection; γ:δ T cells e. involution; thymic stroma. 7–43 A. What is immunological tolerance? B. What is the general name for the antigens against which the immune system is normally tolerant? ANSWERS 7–1 d, e 7–2 d 7–3 b, e 7–4 a
- 13. 9 7–5 d, f 7–6 d 7–7 a 7–8 a, c, d 7–9 c, d, e 7–10 d 7–11 e 7–12 A. Successive gene rearrangement is possible at a TCRβ locus because there are two sets of D, J, and C gene segments downstream of the cluster of V gene segments: (Vβ)n…Dβ1…(Jβ1)n…Cβ1…Dβ2…(Jβ2)n…Cβ2. If a first rearrangement involving Dβ1 and a Jβ1 segment is unproductive, an upstream V gene segment can rearrange to the second D gene segment and an associated J segment. If this is unproductive, no more rearrangements can be made. B. The answer is no. The heavy-chain locus has the following configuration: (V)n- heptamer…23 spacer…nonamer…nonamer…12 spacer… heptamer-(D)n-heptamer…12 spacer…nonamer…nonamer… 23 spacer…heptamer (J)n…Cμ. After the first DJ rearrangement, the intervening D segments between the chosen D and J will be deleted. After VDJ rearrangement, the D segments that lie between the chosen V and DJ will be deleted. Therefore, no unrearranged D segments remain after these two rearrangement events. V and J cannot rearrange directly because the recombination signal sequences are not paired appropriately and do not follow the 23/12 rule; rather, they both contain 23-bp spacers in their recombination signal sequences. Successive gene rearrangement is thus not possible at a heavy-chain locus. 7–13 a—F; b—F; c—F; d—T; e—T 7–14 c 7–15 A. Notch1 is a membrane-bound receptor found on thymocytes that participates in the regulation of early T-cell development. B. Its ligand (Notch ligand) is a membrane-bound protein on the surface of thymic epithelial cells. C. After binding of the extracellular domain of Notch1 to the extracellular portion of Notch ligand, the intracellular domain of Notch1 is released by proteolysis and subsequently translocates to the nucleus. In the thymocyte nucleus, this domain forms a transcription factor complex that displaces repressor proteins from genes involved in T-cell development and initiates transcription of these genes by recruiting transcription activator proteins.
- 14. 10 7–16 c 7–17 a 7–18 a, b, c, d 7–19 a, b, c, e 7–20 b 7–21 c 7–22 e 7–23 b 7–24 a 7–25 A. (i) As well as expressing their own thymus-specific self antigens, medullary epithelial cells in the thymus produce a transcription factor called autoimmune regulator (AIRE), which causes several hundred genes normally expressed in other tissues to be expressed in these cells. The proteins can then be processed to form self peptides that will be presented by MHC class I molecules. (ii) The thymic epithelium uses different proteases for self-protein degradation; cathepsin L is used for peptide production instead of cathepsin S, which is used by other cell types. B. Generating a more comprehensive repertoire of self peptides in the thymus increases the types of potentially autoreactive T cell that are removed from the peripheral T-cell repertoire during negative selection. 7–26 b 7–27 b, d 7–28 a—3; b—5; c—4; d—2; e—1 7–29 a 7–30 d 7–31 The analog of VpreB:λ5 in developing T cells is the protein preTα(pTα), which combines with the T-cell receptor β chain, the first of the two T-cell receptor chains to be expressed, to form the pre-T-cell receptor. The β chain, like the immunoglobulin heavy chain, contains V, D, and J segments. pTα also binds CD3 and ζ components to this complex, and the assembly of the complete complex induces T-cell proliferation and the cessation of rearrangement at the TCRβ
- 15. 11 loci (leading to allelic exclusion). Formation of the analogous pre-B-cell receptor complex of VpreB:λ5 and heavy chain with Igα and Igβ in B cells similarly prevents further rearrangement of the heavy-chain loci. 7–32 e 7–33 d 7–34 A. Only one of the receptors will have to be positively selected for the cell to get the survival signals necessary for it to pass on to the next stage. Even if the other receptor does not react with self MHC this will have no effect on the cell. B. In contrast, both receptors will have to pass the negative selection test for the T cell to survive, because if only one of them fails it, the cell will die. C. Yes. Imagine this situation. The T cell with dual specificity could be activated appropriately during a genuine infection by a professional antigen-presenting cell plus foreign antigen 1 using T-cell receptor 1. But that same T cell, because it is now an activated effector T cell, would also be able to respond to a second peptide, which might be a self peptide, using T- cell receptor 2, without requiring the co-stimulatory signals that only professional antigen- presenting cells deliver. Thus it could cause a reaction against a self tissue, either directly, if it is a CD8 cytotoxic T cell, or indirectly, if it is a CD4 T cell, by activating potentially autoreactive B cells. Furthermore, interferon-γ produced in the response against foreign antigen 1 could activate nonprofessional antigen-presenting cells nearby, inducing the expression of MHC class II with presentation of the self peptide above. Effector T cells with T-cell receptor 2 could make an autoimmune response against it. 7–35 Because T cells drive almost all immune responses, once they have been activated their receptors must continue to recognize the exact complex of foreign antigen and MHC molecule (which does not itself change) that activated them. Because of this requirement for dual recognition (MHC restriction), somatic hypermutation would be more likely than not to change the T-cell receptor to make it unable to recognize either the peptide or the MHC molecule, or the combination of both, thus rendering it unable to give help to B cells or to attack infected cells. This would destroy both the primary immune response and the development of immunity. Even changes that simply increased the affinity of the T cell for its antigen would have no real advantage, because it would not make the immune response any stronger or improve immunological memory in the same way that affinity maturation of B cells does. Also, if somatic hypermutation changed the specificity of the T-cell receptor so that it now recognized a self peptide, this could result in an autoimmune reaction. These considerations do not apply to B cells, because they require T-cell help to produce antibody and will only receive it if their B-cell receptor still recognizes the original antigen. 7–36 A. MHC class II deficiency affects the development of CD4 T cells in the thymus. If the thymic epithelium lacks MHC class II, positive selection of CD4 T cells will not take place. CD8 T cells are not affected because MHC class I expression is unaffected by this defect.
- 16. 12 B. To produce antibody, B cells require T-cell help in the form of cytokines produced by CD4 TH2 cells. Low immunoglobulin levels (hypogammaglobulinemia) in MHC class II deficiency are attributed to the inability of B cells to proliferate and differentiate into plasma cells in the absence of TH2 cytokines. 7–37 A. After thymic atrophy or thymectomy, T cells in the periphery self-renew by cell division and are long lived. B. B cells are short lived and replenish from immature precursors derived from the bone marrow. 7–38 A. Treg suppress the proliferation of naive autoreactive CD4 T cells by secreting inhibitory cytokines. This inhibitory action requires that both the Treg and the other CD4 T cell are interacting with the same antigen-presenting cell. B. Unlike non-regulatory CD4 T cells, Treg express CD25 on the cell surface and the FoxP3 transcriptional repressor protein. 7–39 a 7–40 b 7–41 Rationale: The correct answer is b. Depletion of T cells, but not of B cells, and the absence of a thymic shadow on the X-ray are critical clues. Development of both CD8 and CD4 T cells is affected in this patient because the thymus is the primary lymphoid organ required for T-cell development. Bare lymphocyte syndrome affects either CD8 (type I) or CD4 (type II), but not both. Although patients without a thymus succumb to infections also common in individuals with AIDS, this option can be ruled out because Giulia lacks CD8 T cells, a condition not seen in AIDS patients. Chronic granulomatous disease is a defect of neutrophil, not T-cell, function. 7–42 b 7–43 A. Immunological tolerance is the mechanism that operates to ensure that lymphocytes do not contain antigen receptors specific for host components. This is achieved through a process involving the removal of self-reactive T and B cells called negative selection. Both T and B cells are removed by apoptosis after the engagement of T-cell receptors and immunoglobulins, respectively, if the interaction with their ligands is especially strong. The consequence is the removal of autoreactive lymphocytes that could cause damage to healthy, uninfected cells and tissues. B. Self antigens.
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- 18. rule over Stratovania while I descend to Ohs and take possession of that rich and prosperous country. And, sooner than soon, I will return, bringing you the magic treasure and jewels and the crown and scepter of this Ohsma!" "Oh, but you mustn't!" wailed Jellia, clasping her hands desperately. "Ozma is a real Princess and much more beautiful than I!" "In that case, I shall bring Ohsma back and make her a Starina also!" promised Strut. "Now Hippenscop," he directed, shaking his finger at the odd-looking page, "you and Junnenrump are to obey Jellia in everything. I'll leave three Blowmen here to protect our Starina. The others, and all of my able-bodied fighters, shall fly with me to Ohs." "The Ozoplane holds only four!" cried Jellia, looking desperately over at Nick who was struggling angrily to free himself from the Blowmen. But they had his arms pinioned behind his back, and the poor Tin Woodman was unable to help himself. "Oh, that's all right!" answered Strut, "I and this Tin Emperor will ride in the Friend-ship, and the others will follow on their flying sticks and soon I will return with all the treasures of Ohs!" As the Blowmen started away, shoving Nick and the Soldier ahead of them, Jellia felt so frightened and alone that she burst into tears. "Oh, please, please—couldn't you leave the Soldier to keep me company?" she sobbed, wiping her streaming eyes on her sash. "Of course, if you wish!" Motioning to the Blowmen, they picked up Wantowin as if he had been a sack of potatoes, and tossed him roughly back into the Royal Pavilion. He landed with a clatter at Jellia's feet.
- 19. "But see here! I am not sure I can find the way back to Oz!" protested Nick Chopper as Strut fell into step at his side. "I happened upon this airosphere by the merest chance, and have no idea in which direction Oz now lies." "Just the same—I think you will take me there!" Strut grinned wickedly, tapping Nick on the shoulder with his staff. He already had sent Junnenrump to summon the army, and, glancing over his shoulder, Nick saw a thousand young airmen strutting along behind them. As they came to the shores of Half Moon Lake, Hippenscop came panting and gasping into view. "Her Skyness the new Starina, bade me give you this," he puffed, handing the Tin Woodman the small oil can the Wizard had given him at the party. Nick had forgotten all about his oil can and without it he was likely to rust and become perfectly helpless. Taking it thankfully from the messenger, he hung it on a hook beneath his arm and headed reluctantly for the Oztober. Nick had no intention of flying Strut to the Emerald City. Even if he had to wreck the plane, he would find some way to keep the greedy airman and his legions from conquering Oz. Then he would return and rescue Jellia and the soldier.
- 20. But, without a word to Strut, for argument at this point would have been useless, he mounted the ladder, walked through the cozy cabin and seated himself in the pilot's chair. Strut paused on the top rung of the ladder before he entered. "Follow us closely, men," he commanded gruffly, "no matter how far or fast we fly." Strut's young warriors raised their flying staffs to show that they understood, and with a few final directions, the Stratovanian stepped over the sill, slammed the door of the Oztober and walked rapidly forward, examining everything with lively interest. "So this dragon-body really flys?" he said, bending curiously over the navigator's table. "Ho, what's this? I thought you told me you had no way of finding the route back?" Nick Chopper, much more surprised than Strut, picked up the tidy map that lay on top of the buttons. It certainly had not been there when he left the plane, but here it was now, showing the complete course they had taken since leaving the Emerald City. Concluding this was some of the Wizard's magic, Nick examined the map attentively. Each turn up or down, each mile east or west, was charted accurately.
- 21. "All you have to do is follow this in reverse," exclaimed Strut. Unaccustomed as he was to flying except by staff, he was nevertheless sharp enough to realize the value of a good map when he saw one. "And remember now—no tricks!" he warned, sternly. "Land me safely in Ohs and you will be suitably rewarded. But land me anywhere else and you will be completely obliterated!" Nick said nothing. Weary of Strut's threats and boasts, the Woodman touched the button to inflate the Oztober's balloon, and the "up," "south" and "fast" buttons. In the whirr and splutter of their take-off, the Airlander's further remarks and directions were completely drowned out.
- 22. CHAPTER 9 Jellia in a Frightful Jam For a long time after the departure of the Tin Woodman and of Strut and his legions, Jellia sat forlornly on the Star Throne, trying to stem the tears that coursed slowly down her cheeks. To be stranded on this high and dangerous airosphere was bad enough, but the thought of Strut flying off to destroy Ozma and steal all her treasures was more frightening still. "What on airth shall we do?" questioned Jellia with a rueful smile, of the Soldier with Green Whiskers who was tramping morosely up and down the pavilion. Halting in his march, Wantowin shook his head dubiously.
- 23. "That I cannot say!" he murmured, taking off his cap and staring gloomily inside. "I have no standing in this country at all! But you, Jellia, are a Starina. Therefore you must decide what is to be done. And whatever your Majesty's orders may be, I will carry them out to the letter. To the letter!" declared Wantowin, standing up very straight and tall. "Oh, bother 'my Majesty!'" scolded Jellia. "You know perfectly well I didn't ask to be a Starina of this terrible place!" "It is not what you want but what you are, that counts!" insisted the Soldier, stubbornly. "And there's no getting round it, Jellia, you are a Starina! So while you are deciding what is to be done, I'll just do a bit of reconnoitering. It might be well to know the lay of the air!" "Wait!" cried Jellia as Wantowin started smartly down the steps. "Whatever you do, Wanny—don't run!" she implored earnestly. "You might easily run off the edge and then where'd you be? So do please be careful, and if anything frightens you run straight back here! Do you promise?" "Nothing ever frightens me!" said the Soldier in an offended voice. Marching sternly down the steps he was off at a double-quick, without even a glance over his shoulder. Feeling more alone than ever, Jellia sighed and folded her hands in her lap. But Wantowin's words, foolish as they were, had done her good. After all she was a Starina, for the time being anyway. So, straightening her crown, and drying her tears, Jellia tried to think how she should act under such bewildering circumstances. How would Ozma act, for instance, if she were sitting on the throne of this singular airtry? Even thinking of the gentle and dignified little Girl Ruler of Oz, steadied Jellia. Holding her head very high, she stepped down from the dais and began pacing slowly up and down the pavilion, switching her green skirts in such a regal manner that the two messengers who had returned quietly to their posts, stared at her with new interest and admiration.
- 24. "Is there anything we might bring your Strajesty?" asked Junnenrump, bowing from the waist and clicking his heels smartly together. At his question Jellia paused and eyed the two, speculatively. "Why, yes," she decided after a moment's thought. "You, Junnenrump, may send some one to amuse me, and you, Hippenscop, may bring me two of those winged staffs. It is neither safe nor proper for a Starina and her Army to be without them!" "But, your Skyness!" Hippenscop leaped into the air and spun round and round in an agony of embarrassment. "There are no extra staffs!" he blurted, finally coming to a stop before her. The little fellow looked so distressed, Jellia was on the point of letting him off. Then, remembering just in time that she was bound to be obeyed, she raised her arm. "Go!" she commanded haughtily. "And do not return without two winged staffs!" Junnenrump already had started, and at Jellia's stern command Hippenscop backed dejectedly down the steps, his eyes bulging with dismay and consternation.
- 25. "If Wanny and I had flying sticks, we'd at least be as well off as the rest of these Airlanders," reasoned Jellia, resuming her walk. "But what funny names," she mused, as the messengers disappeared in two different directions and at two different speeds. "They make me think of—" here Jellia took a little run and jump, following it with a skip and a hop. "I suppose" she continued, talking conversationally to herself, "that is what their names really mean, everything is so mixed up here." Regaining her throne in one long slide, Jellia brought up with a slight start. This, she decided, was no way for a Starina to act. Smoothing down her dress, she walked sedately to Strut's throne and reached underneath. The real reason she had got rid of the messengers, of course, was so she could recover the kit- bag and have a chance to examine its contents without being observed. The cheer gas had saved them on one occasion, and perhaps there was magic powerful enough to enable her and the Soldier to escape from the airosphere before Strut returned. The bag was still there and snatching it up in her arms, Jellia climbed back on the throne. But just as she was about to zip it open, Junnenrump bounded up the steps of the pavilion, dragging a lean old Skylander by the hand.
- 26. "His Majesty's Piper!" announced Junnenrump, giving the Piper a shove forward and seating himself expectantly on the messenger's bench. Jellia was annoyed to have Junnenrump return so soon. But since she had sent for someone to amuse her, she could not very well object. So, resting her chin in her hand she looked curiously at the royal Piper. The old Skylander was tremendously tall and thin. His tunic was short and plaited, and under his arms he carried a pair of enormous bag pipes. Jellia never had cared for bag pipes, but on an airosphere she supposed wind instruments such as this naturally would be popular. The Piper, however, did not immediately play on his pipes. Instead he struck a few light and pleasant chords on the top buttons of his tunic. "Shall I do a buck and wing, or a little Skyland fling? Shall I sing a little sing, for you, Dear?" bawled the Piper cheerfully. He looked so funny that Jellia burst out laughing. Thus encouraged, the Piper proceeded to sing, punctuating his song with extraordinary leaps and toe tappings. "When we Skylanders feel low, we just Dance the stratispho; Step it high, kick and fly, toss the Partner up ski-high. High HO! "Would you care to try it?" he asked politely, holding out his hand to Jellia. "No, No! Not today!" gasped the Oz maid, backing as far as the star throne would allow. "But I've really enjoyed watching you very much, and your singing is lovely," she added, generously. "Ah, but wait until you hear me play," puffed the Piper. Raising his pipes he blew forth such a hurricane of whistles, squeals and fierce thunderings that poor Jellia clapped both hands to her ears. "Tell him to go away," she screamed above the awful din, wildly motioning to Junnenrump who was tapping his foot in time to the
- 27. pipes and looking highly diverted. "Tell him to come back tomorrow." The fierce music of the bag pipes had brought airlanders running from every direction. Crowding round the pavilion they waved and bowed to the new Starina. Realizing she never would have any privacy under the Imperial Canopy, Jellia slipped off her throne. The messenger had the Piper by the tunic tails and was easing him gently down the steps. Jellia waited till they reached the bottom, then, as all the airlanders began to run after the still furiously pumping piper, Jellia started in the opposite direction. Surely somewhere, she thought, clutching the kit-bag close to her, somewhere she could find a quiet corner or cave or clump of bushes where she could examine the contents of the Wizard's bag without interruption. So anxious was Jellia to be by herself, she broke into a run. Failing to notice a crystal bar stretched across the path, she tripped and fell violently up a tune tree. Falling down is bad enough, but falling up is worse still. Jellia not only had barked her shins on the crystal bar, but had bounced into the air so high she lost her breath and plunged down so abruptly among the top branches of the tune tree that she
- 28. was somewhat scratched and shaken. She knew it must be a tune tree because plump black notes grew in clusters like cherries between the leaves. Several, dislodged by her fall, broke into gay little arias and chords. At any other time Jellia would have been quite interested, but now she was too agitated and upset to care. "Such a country—or airtry!" groaned the Oz maid, rubbing her left ankle and her right knee. "One can't even fall down in their own way!" Parting the branches the ruffled little girl looked crossly out. It was quite a long way to the ground, but nevertheless Jellia decided to climb down. But suddenly it occurred to her that the top of the tune tree was as good a place as any, to open the kit-bag. Easing herself to a larger limb, she balanced the bag carefully in her lap and stretched out her hand to pull the tail. Then a piercing scream and the thump of a hundred footsteps made her draw it back in a hurry. Parting the branches of the tree for a second time, she saw Wantowin Battles running toward her like the wind. "Help! Help! Save me!" yelled the Soldier with Green Whiskers. And he had reason to yell for just two leaps behind him panted Kabebe, waving an enormous crystal rolling-pin. After the Queen pounded the three big Blowmen, and after the Blowmen came nearly a hundred men, women, and children. Before Jellia had time even to guess why they were chasing the Army, Wantowin tripped over the same crystal bar that had caused her upfall, and landed with a terrific grunt in the branches beside her, scattering half and quarter- notes in every direction. The Airlanders stopped short and watched with breathless interest as the Soldier disappeared into the thick foliage of the tune tree.
- 29. "What's the matter? What happened?" whispered Jellia reaching out to steady the soldier who was bouncing wildly up and down on a nearby limb. "YOU?" gasped Wantowin, almost losing his balance at the shock of seeing her. "Oh, Jellia! We must leave at once! At ONCE! As I was passing the cooking caves, Kabebe rushed out and grabbed me. She has decided to blow us away most any minute now. She has persuaded the Airlanders that Strut is lost and never will return. Oh why, WHY, did we ever fly to this terrible place?" "Be quiet!" hissed Jellia, frightened almost out of her wits at this new turn of affairs. "How can I think with you making all that noise?" "Come down! Come down!" bawled Kabebe. "Come down before I shake you down!" Grasping the trunk of the tune tree she gave it a playful shake. Rolling his eyes up, the Soldier glanced desperately at Jellia, and Jellia, as desperately, glanced back. "You might as well go down," she whispered resignedly, as the Queen gave the tree a tremendous shake that nearly dislodged them
- 30. both. "Not without you," shivered Wantowin, hugging his branch for dear life. "Oh, well—let's get it over with," said Jellia despairingly. "Blowing away may not be so bad, and I'd rather do anything than stay up here." Tucking the kit-bag under one arm, Jellia swung herself down by the other and dropped lightly to the ground. "What is the meaning of this outrageous behavior?" she demanded, as Wantowin dropped fearfully beside her. "His Majesty shall hear of this, I promise you!" Kabebe, astonished to see Jellia as well as the Soldier with Green Whiskers drop out of the tree, took a hasty step backward. Jellia quickly followed up her advantage. "I'm amazed!" she said sternly. "I thought you knew that I was to help you rule while King Strut is away!" At this bold speech, Wantowin looked at Jellia in round-eyed admiration. Though her cheeks were scratched and her crown slightly askew, the little Waiting Maid looked every inch a ruler's helper, if not a ruler. Even the Blowmen began to shift uneasily from one foot to the other, their mouths falling open at Jellia's indignation. But Kabebe raised both arms and fairly screeched at the little Oz Maid. "How dare you speak to me like that?" she shrieked. "King Strut is lost and never will return! I am Queen here—and I don't need your help! Blowmen! Seize this impudent pair, march them to the edge of the cliffs and blow them away." The crowd of Stratovanians looked uncertainly from Kabebe to Jellia. "His Highness left you here to protect me!" Jellia reminded them sternly. But even as she spoke, she knew they had decided to obey Kabebe. She was flashing her star eyes so threateningly, and waving her winged stick so close to their heads, that the Blowmen were afraid to defy her.
- 31. "Come along, now," grumbled the first Blowman, taking Jellia roughly by the arm. "You've made enough trouble here!" The other two Blowmen seized the trembling Soldier and began marching sternly toward the edge of Strut's Skyland. Jellia pulled back with all her strength, as also did Wantowin, but, hustled along by the huge Skylanders, they could do little to help themselves. Relentlessly, with the jeering citizens of Stratovania running along after them, the unfortunate Oz pair was dragged on. "Just wait till your Master hears about this," sobbed Jellia, as the Blowmen shoved them as near to the edge of the cliffs as they dared go themselves. Then they stepped back to lift their horns. Jellia had managed to retain her hold on the Wizard's kit-bag, but even so she felt that their last moment had come.
- 32. Jellia gave a final sad little wave to the Soldier, who really was quite brave now that his doom had arrived. The Blowmen pointed their horns straight at them, but before they even could inflate their cheeks, a fierce roar and splutter from the clouds caused every head to turn upwards. "The ship—the ship! The flying ship!" cried the First Blowman, letting his horn fall disregarded to the ground.
- 33. "It's Strut!" screamed the Stratovanians, treading on one another's toes in their sudden frenzy to be out of sight of their Master when he landed. "'Tis the Master himself!" cried the first Blowman, yanking Jellia and the Soldier back from the edge of the Skyland. Pulling Kabebe along with them, the Blowmen ran as never before, closely followed by Strut's scurrying subjects. One moment later there was not a single airbody in sight. Convinced that their cruel and brilliant ruler had returned, they ran like rabbits. Some even flew, helping themselves along with their winged staffs, while Jellia, sinking on a large, crystal boulder, stared dazedly at the silver-bodied plane dropping rapidly toward them. "It can't be the Oztober!" cried Jellia, delightedly. "It couldn't have come back so soon!" "It's not!" cried Wantowin Battles, tossing up his cap and waving his arms exuberantly. "It's the other one, the Ozpril, and that means—" In his extreme excitement, the Soldier tripped over a balloon bush and fell seven feet into the air. "It means the Wizard himself has come to help us," sputtered Wantowin, blinking rapidly as he landed
- 34. hard on the rock beside the young Oz maid. "Three cheers, Jellia! The Wizard of Oz has saved us!"
- 35. CHAPTER 10 The Wizard in Stratovania! It was indeed the Ozpril, just as the Soldier with Green Whiskers had said. Even at a distance, Jellia could spell out the name on the gleaming body and, as the silvery plane came swooping toward them, she could not repress a shout of joy. Too exhausted by the dreadful ordeal she had just been through to run to meet the ship, she jerked off her scarf and waved it wildly over her head. About ten feet from the crystal boulder on which she had been sitting, the Ozpril came to a gentle and perfect landing. Scarcely had the whirr and sputter of its engine died away before the door of the cabin burst open and down climbed the little Wizard of Oz, followed by Dorothy and the Scarecrow. The Cowardly Lion, last of all, had difficulty fitting his paws on the rungs and, after a trembling descent, rolled over on his back, his four feet straight up in the air. The trip had not agreed with the Cowardly Lion at all. Weak and dizzy, he made no attempt to rise.
- 36. "Here you are at last!" cried the Wizard happily, rushing over to Jellia and seizing both of her hands. "So THIS is where you've been! Well I must say it's a fine place. Why it's beautiful, beautiful!" Swinging round so he could look in all directions, the Wizard positively glowed with interest and enthusiasm. "What's so beautiful about it?" growled the lion without turning over. "Is there any grass? Are there any trees? Is there anything to eat?" Dorothy, on the point of embracing Jellia, gave a little scream, for the Tell-all-escope, which she had picked up just before leaving the plane, was making terse announcements. At this point it happened to be pointed at Jellia. Clearing its throat it remarked in a superior way: "You are now looking at Miss Jellia Jam, formerly of Oz, at present new Starina of the Strat, by edict of Strutoovious the Seventh. Miss Jellia Jam, Starina of Stratovania! Period! Stop, drop or point elsewhere!" "Why, Jellia!" gasped Dorothy, letting the Tell-all-escope fall with a crash, "are you, really? Oh my! I don't suppose you'll ever want to return to Oz, now. Why, you must be having a wonderful time!"
- 37. "Humph!" sniffed Jellia, with a slightly wan smile. "If being pinched, chased and nearly blown to atoms is having a wonderful time, then I guess I've been having it all right!" "Tell me," requested the Scarecrow, who had been walking in a slow circle around Jellia. "Does one prostrate oneself before a Starina, or does one merely kiss her hand?" "Neither," laughed Jellia. Jumping up she gave the Scarecrow such a hug he was out of shape for hours. "But quick!—Let's all hop in the Ozpril and fly away before something terrible happens." "Fly away?" cried the Wizard, shoving back his high hat. "But, my dear—we've only just come! I've been flying all night and need a little rest and refreshment before we start off again. Besides, I would like to see more of this interesting airland and its people, and add to my data on the Strata." "That's what Nick thought," observed Jellia, putting both hands on her hips. "And look what happened to him!"
- 38. "What did happen to him?" demanded the Wizard, realizing for the first time that Nick was not among those present. "You tell him," sighed Jellia to the Soldier. Sinking back on the boulder she held her aching head in both hands. All eyes turned toward the Soldier with Green Whiskers who opened and closed his mouth several times without saying a word. The Wizard, now thoroughly alarmed, began shaking him on one side and the Scarecrow on the other, until finally Wantowin took a tremendous swallow and gave them the whole story. When the narrator reached the part where Strut had ordered Nick and him blown away, the Scarecrow hurried over to the balloon bush and began picking the almost ripe balloons as fast as his clumsy cotton fingers would permit. Not till he had about twenty did he even pause. So light and flimsy was the straw man that the bunch of balloons on their long stems kept jerking him into the air. After each jerk he would give a little grunt of satisfaction. "These are just to keep me aloft—in case of accidents," he explained hastily to Dorothy who was watching him intently.
- 39. "But what of us?" asked the little girl, looking anxiously toward the Canopied City which, at present, seemed absolutely deserted. "You say that this wretched Strut, after naming Jellia Starina, forced Nick to fly him to Oz?" exclaimed the Wizard, grasping Wantowin Battles by both arms and gazing into his face. "Not only that," Wantowin told him hoarsely, "but he's taken his Blowmen and a thousand fighting men to conquer the country! He intends to bring back Ozma's crown, scepter, jewels and all the treasures in our castle!" finished the Soldier, dolefully. "Oh, can't we do something Wizard?" cried Jellia determinedly. "I simply won't be Starina! I won't! I WON'T!" "Just the same—you make a very pretty one," murmured the Scarecrow, patting the little Oz Maid consolingly on the shoulder. "But of course, we cannot allow this bounding airlander to take Oz!" "If Nick had not 'taken possession' of Stratovania for Ozma, he'd never have thought of it," groaned Jellia. Rising stiffly, she picked up the kit-bag from the crystal rock beside her. "Ah—so you still have my magic kitty!" In spite of his anxiety the Wizard smiled. "Indeed I have," said Jellia firmly. "It saved us from being blown away. I used some of your cheer gas, Wiz, but I didn't have time to try out any of the other magic. Here, you'd better take it now and do let's be starting. No telling when Kabebe and those three Blowmen will be coming back." "Forward march! Forward march!" Wantowin Battles started off all by himself for the Ozpril. "Hurry, hurry!" he called over his shoulder. "If those fearful people return they'll surely make trouble!" yelled the Soldier, his voice growing more emphatic. "Well, it's certainly a mix-up," said Dorothy, moving closer to the Wizard.
- 40. "What do these people look like, Jellia?" she asked curiously. "Really I'd enjoy seeing a few." "They look like nothing you ever have imagined!" Jellia told her with a slight shudder. "Goochers! Here come some now! And oh—it's those Blowmen—and all the others! Look, Wizard! Could we reach the Ozpril before they reach us?" "Let's not try," decided the Wizard, as the Blowmen broke into a run. "Even if we made the plane, they might blow us to bits before I could get her started. Let's stay here and reason with them till I find something in this bag to help us." "Oh, woe is we! Oh, woe is we!" gulped the Scarecrow, taking little runs and leaps into the air, hopeful that his balloons would lift him out of the danger zone as the threatening company drew closer. The Queen was marching grimly ahead of her subjects. In some way, decided Jellia, she had discovered Strut had not been in the silver plane. As the Wizard opened his kit bag the little Oz Maid rushed over to the Cowardly Lion.
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